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PROCEEDINGS

OF THE
GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

OF THE

ZQVOLOGICAL SOCIETY

OF LONDON.

1911, pp. 557-1213,

witH 48 Puatres and 86 TEx-FIGURES.

LLL585 |
PRINTED FOR THE SOCIETY,
AND SOLD AT THEIR HOUSE IN REGENT’S PARK.
LONDCN:

MESSRS. LONGMANS, GREEN, AND CO,
PATERNOSTER ROW.

Ss a

OF THE

COUNCIL AND OFFICERS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.

WON Te

COUNCIL.

His Grace Tut Duxke or Beprorp, K.G., President.

Tue Karu or Auramont, F.S.A.

Sir Joun Rose BRADFORD,
KC MEGS) MED EDS,
FE.R.S., Vice-President.

Lt.-Col. Srp R. HaAvetock-
Cuartes, K.C.V.O., M.D.

ALFRED H. Cocks, Esq., M.A.

The Rt. Hon. the Ear or
@rommr, | 2-0 1G-C bs
G.C.M.G.

F. G. Dawtrey Drewirt, Ksq.,
WieA\o5 WilslD).

CuarLtes DrumMonn,
Treasurer.

Str Epwarp Duranp, Bt., C.B.

FREDERICK GILLETT, Esq., Vice-
President.

Srpney F. Harmer, Ksq., M.A.,
Se.D., F.R.S., Vice-President.

Ksq.,

Sir WALTER Roper LAWRENCE,
Bt. G.C.AnK-

Sir Epmunp G. Lopmr, Bt.

EK. G. B. Meapre-WaAtpo, Esq.,
Vice-President.

P. Cuaumers Mircuet, Esq.,
MEARS UDESc:.) JElonas ae
E.B.S., Secretary.

W. R. Ocitvie-Grant, Esq.

ADRIAN D. W. Poutock, Esq.

A. Trevor- Battys, Ksgq.,
M.A.

AntHony. H.
Ksa.

A. SmirH Woopwarp, Ksq.,
LL.D.,F.R.S., Vice-President.

Henry Woopwarb, Hsq., LL.D.,
F.R.S., Vice-President.

WINGFIELD,

PRINCIPAL OFFICERS.
P. CHatmers Mircnety, M.A., D.Sc., Hon.LL.D., F.B.S.,

Secretary.

Frank HK. Bepparp, M.A., F.R.S., Prosector.

R. I. Pocock, F.R.S., F.L.S., Curator of Mammals, and
Resident Superintendent of the Gardens.

D. Sera-Suiru, Curator of Birds and Inspector of Works.

Epwarp G. Boutencer, Curator of Reptiles.

Henry G. Purmimer, F.R.S., M.R.C.S., Pathologist.

F. H. Waternoust, Librarian.

JOHN Barrow, Accountant.
W. H. Coun, Chief Clerk.

dy?

LIST OF CONTENTS.

EXHIBITIONS AND NOTICES.

Page
The Secretary. Report on the Additions to the Society's
Menagerie during the month of February 1911 ......... 557

Mr. D. Sern-Sairs, F.Z.S. Exhibition of a living Hybrid
between the White-eyed Pochard (Aythya nyroca) and
the Marbled Duck (Marmaronetta angustirostris) ...... 558
Mr. R. 1. Pocock, F.RS., F.L.S., F.Z:S. -Exhibition of
the skin of a new Chacma Baboon (Papio porearius,
subsp. griseipes), a specimen of the North American
Black-footed Polecat (Putorius nigripes), ete. (‘Texti-

Sir E. Ray Lanxestsr, K.C.B., F.R.S., F.Z.8. Exhibition
of a special supplement to the ‘ Field’ newspaper ...... 620

Dr. R. T. Lererr, F.Z.8. A demonstration of Nematode
parasites obtained from animals in the Zoological
Garden sy hk eh isdn Wea ee. cae SpE AA EE 620

Mr. R. I. Pocock, F.R.S., F.L.8., F.Z.S. Exhibition of
a newly born Masked Palm-Civet (Paradoxurus larva-
tus), with an abnormal leg. (Text-fig. 147.) ............ 621

The Secretary. A letter from the Governor of the Seychelles
on Wand=Mortoises 1m those Islands 2.0.2... .........4....- 622

Dr. H. B. Fanruam, F.Z.S., and Miss Annie Porter, D.Sc.
On a Bee-disease due to a Protozoal Parasite (Vosema

1V

The Secretary. Report on the Additions to the Society’s
Menagerie during the month of March 1911 ............

Mr. C. Tare Reean, M.A., F.Z.S. Exhibition of a series of
lantern-slides of scales of the Salmon (Salmo salar) ...

Mr. D. Seru-Suiry, F.Z.S. Exhibition of a nest of the
Grey Struthidea or Apostle Bird (Struthidea cinerea),
and lantern-slides of Penguins in moult and of wild
Swainson’s Lorikeets (Zvrichoglossus nove-hollandic).
(EGS NGS oP ern Ahi oS Seno udavadonasudoasesG

Mr. C. Curisry, F.Z.S. Exhibition of a collection of skins
from Uganda and of a loin-cloth taken from a native
Tow Northern Nieeria: ..ceeceeneerere had cena nee

Dr. Wit11AM Nico, M.A., F.Z.8. On a unique Patholo-
gical Condition ina Hare. (Text-fig. 165.) ............
Mr. R. I. Pocock, F.R.S., F.LS., F.Z.8. Exhibition of
some hair from the “puppy coat” of a Grey Seal
(Halichcerws giiypus) a eseee ace ene eee eRe ee

Mr. E. G. Boutencer. Exhibition of living male specimens
of the Midwife Toad (Alytes obstetricans) carrying the

COGS coerce cr cece cece eee ee rece eee cer eee nesessreecr scenes ceeeeene

Mr. A. E. Anperson. Exhibition of photographs of fossil
mammals in the American Museum of Natural History .

The Secretary. Report on the Additions to the Society’s
Menagerie during the month of April 1911 ............

The Rey. A. Mines Moss, M.A., F.Z.S., F.E.S. Notice of a
Mmemomon the Sphingides oll eru case eee eeere ee eeeee

Page

671

671

671

674
696
696
696

869

869

Mr. H. G. Purmmer, F.R.S. Report on the Pathological .

Examination of Rats (Jus decumanus) caught in the
Regent's Park and in the Society’s Gardens

eroesccecens

Dr. R. W. Ssurexipr, C.M.Z.S. Exhibition of a photograph
of a male albino Woodchuck (Arctomys POMTAS)) saecce a0:

Mr. R. E. Hotpine. Exhibition of the Horns of a High-
land Ram, a Fallow Deer, and a Roebuck, which were
fused at the base, and the skull of a coursing Grey-
hound with abnormal dentition

Seer eee eecces sesso ceo essere

Vv
Page
Mr. R. I. Pococr, F.R.S., F.L.S., F.Z.S. Exhibition of, and
remarks upon, the skin and skull of a specimen of the
Crested Rat (Lophiomys ibeanus Thos.). (Text-fig. 190.) 946

The Secretary. Report on the Additions to the Society’s
Menagerie during the month of May 1911 ............... 985

Mr. D. Sera-Suirn, F.Z.8. Exhibition of, and remarks
upon, two immature Black-backed Porphyrios (Por-
phyrio melanonotus) with wing-claws. (Text-fig. 200.) 985

Mr. J. Lewis Bonuots, M.A., F.L.S., F.Z.S. Exhibition of
a pair of abnormally coloured Egyptian Desert-Mice
(A GPUOTMES GRUSSOIS)) REN Roe nORe eos Ane odie soe danneteeee st 986

Dr. W. T. Catman, F.Z.8. Exhibition of living specimens of
the Brine Shrimp (Artemia salina), bred from Tidman’s
SOa Calc eee tet mem ace ae teres eee reat etea te tc cecal te erases centre 986

The Secretary. Remarks upon the eggs of Struthio massai-
' cus, S. australis, and S. molybdophanes, seen at Mr. Carl
Hagenbeck’s Ostrich Farm at Stellingen.................. 987

The Secrerary. Remarks upon a pair of young African
Rhinoceros seen at Mr. Carl Hagenbeck’s Zoological
JEpue enn tee) MEU. 0 roan Jane AB ac GAR AE AAlNBn noth eb 987

Mr. R. I. Pococn, F.R.S., F.L.S., F.Z.S. Exhibition of
photographs of, and remarks upon, two hybrid foals,
Equus asinus somaliensis x Hquus zebra and Hquus
asinus somaliensis x Hquus quagga chapmanni. (Text-
{Sea AD pears epee oman dee c aes aucei md atea eae ems ocibh 988

PAPERS.

26. On the Amphipod Genus Leptocheirus. By EK. W.
Sexton, Marine Biological Laboratory, Plymouth.
(Elates Revell SXGRXe and Wextoties WAG ee sss. ce anna cee 561

27. Notes on Marine Ostracoda from Madeira. By G.
STEWARDSON Brapy, M.D., LL.D., D.Se., F.R.S.,
OpWilAds. (IPI Nias OCC OS IUD) ei hansoccs sag sbnen sean Contes 595

28.

30.

ol.

33.

34.

36.

vi

On Colour and Colour-pattern Inheritance in Pigeons.
By J. Lewis Bonuors, M.A., F.L.S., F.Z.8., and F. W.
Sud pune H ZS. ua(Plaibes 2OXUuEL —XOXGV cl) I Eee ee

. Contributions to the Anatomy and Systematic Arrange-

ment of the Cestoidea.—I. On some Mammalian
Cestoidea. By Frank E. Bepparp, M.A., F.R.S.,
F.Z.8., Prosector to the Society. (Text-figs. 148-159.).

On the Natural History of Whalebone Whales. By
J. A. Morcu, Christiania, (Text-figs. 160-163.) ......

On Three New Trematodes from Reptiles. By WiLLrAm
Nicott, M.A., D.Sc., M.B., F.Z.S. (Plates X XVII.
& XXVIII.) |

Coe eee pret ese eres ees e terse e trees sere ee essere oesees

. The Duke of Bedford’s Zoological Exploration of Hastern

Asia.— XIV. On Mammals from Southern Shen-si,
Central China. By Otprienp Tomas, F.R.S., F.Z.S.
(Plate X XIX.)

freee ees e weer sere esr ese ese esse eseeeresesseseves

An Investigation into the Validity of Miillerian and
other Forms of Mimicry, with Special Reference to
the Islands of Bourbon, Mauritius, and Ceylon. By
Nevint—E Manpers, Lieut.-Colonel, R.A.M.C., F.Z.S.,

CSCC ee eee mw ene re reer essere seseseeessesserersesessessesesseoes

The Distribution of the Avian Genus I/egapodius in the
Pacific Islands. By J.J. Lisrer, M.A., F.R.S., F.L.S.,
OAS. ai Mexctiatigs NGG:)iss nC ie BER ene Renny aon Renvee

. Contributions to the Morphology of the Group Neritacea

of the Aspidobranch Gastropods.—Part Il. The
Heticinip#, By Ginpert C, Bourne, M.A., D.Sc.,
F.RS., F.Z.8. (Plates DOK I), Reese

On the Palatability of some British Insects, with Notes
on the Significance of Mimetic Resemblances. By
R, I. Pocock, F.R.S., F.LS., F.Z.S., Superintendent
of the Society's Gardens and Curator of Mammals.
With Notes upon the Experiments by Prof. E. B.
Poutton, F.R.S8., F.Z.S.

Coe eae eee e eee cesesesoeseevsceecsevere

. The Aleyonaria of the Cape of Good Hope and Natal.—

Gorconacea. By J. Stuarr THomson, Ph.D., F.R.S.E.,
F.LS., Lecturer and Senior Demonstrator in Zoology,
University of Manchester. (Plates X LIII.—_XLV. and
Wext-He, «NG7,).ccccin kee stes sates te ees nas LT eGeaee

Page

601

626

677

696

749

759

809

870

38.

39.

40.

4].

43.

44,

47.

Vil

On the Structure of the Skull in Cynodont Reptiles.
By R. Broom, M.D., D.Sc.,-C.M.Z.S. (Plate XLVI.
and Text-figs. 168-180.)

ew ree meee eee eee eee e erst ee sesesserere

Tooth-germs in the Wallaby (Macropus billardieri). By
A. Hoprweti-SmityH, L.R.C.P., M.R.C.S., and H. W.
VAREn iss) Moa MODE hase shEZS. (Plate
DIG Ls ewavch Mescpsintets pales | alCl)) a4. bad saceash ab seoeuaaeeee

On a New Species of Dinotherium (Dinotherium hobleyi)
from British East Africa. By C. W. AnpreEws, D.Sc.,
F.R.S., F.Z.S. (British Museum, Natural History).
(RP Tasbe XoTEIVATSIMIE esse 2 casts Narcuiestctinsiu es sic ae eersamelaticticl ae a

On an Amphipod from the Transvaal. By the Hon.
Pau A. MerHuen, F.Z.8. (Plates XLIX.—LI.) ......

. An African Rhinoceros, Klipspringer, and Gazelle. By

R. Lypexker. (Text-figs. 191-193.)

weet eee eee see tesco wee

The Subspecies of the Spanish Ibex. By Prof. ANGEL
Caprera, C.M.Z.S. (Uelaues 1 ye —LIV. and Text-figs.
194— 199.) EN ata clits Ae RR ama Pe Sa lal a Re

On Antelopes of the Genera Madoqua and Rhynchotragus
found in Somaliland. By R. E. Draxs-Brockmay,
IME ChS Wak C le) EeZeS (Plates 1a\Viids IVils)en

. Contributions to the Anatomy and Systematic Arrange-

ment of the Cestoidea.—II. On Two New Genera of
Cestodes from Mammals. By Frank E. Bepparp,
M.A., F.R.S., F.Z.8., Prosector to the Society.
(Text-figs. 204-215.)

. Some Madreporaria from the Persian Gulf. By Rurn

Harrison, Oxford. With a Note on the Memoir and
some Further Notes on Pyrophyllia inflata by SYDNEY
J. Hickson, M.A., D.Se, F.R.S., F.Z.8. (Plates
VIL. & LUVIN. and Text-figs. 216—221.)...............

On Variation in the Medusa of Merisia lyonsi. By
CHARLES L. Boutencrer, M.A., F.Z.S., Lecturer on
Zoology in the University of Birmingham. (Plate
EXG en eM erate SD PE 228. esc ciwia tcl so ciewesiainine oui

Page

893

958

963

977

994

1018

Vill

Page
48. The Marginal Processes of Lamellibranch Shells. By
Cyrit CrossLanb, F.Z.S. (Plate LX. and Text-figs.
IOUS gl hE Mecctle cs ieiteiais inte wiet:o Sell eieitine ae «eae tae iS oer 1057
49. Warning Coloration in a Nudibranch Molluse and in a
Chameleon. By Cyrit CrossuanD, F.Z.S. ............ 1062
50. Chromodorids from the Red Sea, collected and figured
by Mr. Cyrin CrossuanD. By Sir CHAries E ior,
TEAC INING Io, Opleen Ia Ass (Celene IX.) «occossas0a0en00c 1068

51. On some New South African Permian Reptiles. By
R. Broom, D.Sc.,C.M.Z.S. (Plates LXII. & LXIIT.). 1073

52. On a New Tree-Frog from Trinidad, living in the
Society's Gardens. By Epwarp G. BouLENGErR,
(Chennai Ci Iaveroalles” (leleiws, IDOI 3) soncaeonssocoscn6 1082

53. A Contribution to the Ornithology of Western Colombia.
By C. EK. Hetimayr, Division of Birds, Zoological
MuseunosMumich’ aa. cere ec eeerer cece reece rer er eee reenter 1084

ADDENDUM.

Additional reference to Dr. P. CHatmers MircHe.’s
memo, “On Longevity and Relative Viability in
Mammals tamdaltirds'? 2) Sete Saeeaaie oe ene 868

siprateiesd cbratslat avelue | Srtais AES ee ae Eee ee XVI1ll
indexok Scrembitie Names s.45.-)) nee eee ee X1X

siseisar ang aomaseee Resco eee ae eee LEE XXXVIil1

AU RSIVAG BS MN CYAN pasar

OF TIIK

COND Wye Oks;

With References to the several Articles contributed by each.

1911, pp. 557-1218.
2s

Anpprson, A. EK.

Exhibition of photographs of fossil mammals in the
American Museum of Natural History

ANDREWS, CHARLES W., D.Sc., F.R.S., F.Z.8. (British
Museum, Natural History).

On a New Species of Dinotherium (Dinotherium
hobleyt) from British East Africa. (Plate XLVIII.)

Bepparb, Frank E., M.A., F.R.S., F.Z.S., Prosector to the
Society.

Contributions to the Anatomy and Systematic Arrange-
ment of the Cestoidea.—I. On some Mammalian
Cestodeawn Hexboiess U4 Salo) imme ttm ite ane vacultns

696

943

626

x
Page
BEDDARD, Frank E., M.A., F.R.S., F.Z.8. (Continwed.)

Contributions to the Anatomy and Systematic Arrange-
ment of the Cestoidea—II. On Two New Genera of
Cestodes from Mammals. (Text-figs. 204-215.) ......... 994

Bonnore, J. Lewis, M.A., F.LS., F.Z:8.

Exhibition of a pair of abnormally coloured Egyptian
ID eseitin Wess (CUI IOIES GRUSSUIS) \sosascongsaaddoscoba5snonanAsosor 986

Bonnore, J. Lewis, M.A., F.L8., F.Z.S., and Smatiey,
FREDERIC W., F.Z.S.

On Colour and Colour-pattern Inheritance in Pigeons.

(Bilates XXII XC NVI ie ete nega Ne er hese ee ee 601

BouLeNnGer, CHARLES L., M.A., F.Z.S., Lecturer on Zoology

in the University of Birmingham.

On Variation in the Medusa of Merisia lyonsi.
(elenng JOD ane INex giles, WY2= 2) sn acadonoascconnancoce 50 1045

BouLencer, Epwarp G., F.Z.S., Curator of Reptiles.

Exhibition of living male specimens of the Midwife
Toad (Alytes obstetricans) carrying the eg@s............000666 696

On a New Tree-Frog from Trinidad, living in the

SoCs Ceiling, . Celis GIDC) sdohannonsscsocncnoaoscnaes 1082

Bourne, GinBert C., M.A., D.Sc., F.R.S., F.Z.8.

Contributions to the Morphology of the Group Neritacea
of the Aspidobranch Gastropods.—Part II. The Huts-
cinipm. (Plates XXX—XUIL.) w..n.0.. Bt ogas AOR 759

x1

Page
Brapy,- G. Strewarpson, M.D., LL.D., D.Se., F.BS.,
C.M.Z.S.
Notes on Marine Ostracoda from Madeira. (Plates
DOGO). Seladiged ARORA? BERN aH See asa RMA ae ayaa Rin 595
Brockman, Raupy E. Drake-. See Draky-Brockman, R. E.
Broom, Rosert, M.D., D.Sc., C.M.Z.8.
On the Structure of the Skull in Cynodont Reptiles.
(late Xavi and Wext- figs hGS G05 ioe rcueeeaensee ans. 893
On some New South African Permian Reptiles.
(Elenite Bp Ue ca bp. WEE) tence bie: Renate bear cbadtanadnceosnac 10735
Caprera, Prof. ANcrn, C.M.Z.S.
The Subspecies of the Spanish Ibex. (Plates LII.—
JULI, uae ARexeiaitresi Ie ES Ie) eecna ge aeetiadade ssoubeboseokace: 963
CALMAN, Wii ho IDEST@.5 1GZelSy
Exhibition of living specimens of the Brine Shrimp
(Artemia salina), bred from 'Tidman’s Sea Salt ............ 986

Curisty, Curapert, M.B., F.Z.8.

Exhibition of a collection of skins from Uganda and

of a loin-cloth taken from a native in Northern Nigeria. 672

CRossLAND, Cyrit, F.Z.S.

The Marginal Processes of Lamellibranch Shells.
(Gelenie) IDG, eumel WMasxncsieys, P79) 5 A310), \eacemcandacodandde sobeae 1057

Warning Coloration in a Nudibranch Mollusc and in
a Chameleoignemey tytn cue tac n Ment Atk oot ecane 1062

Xi
Draxe-Brockman, Raupu E., M.R.C.S., L.R.C.P., F.Z.8,

On Antelopes of the Genera Madoqua and Rhyncho-
tragus found in Somaliland. (Plates LV. & LVI.) ......

Exsot, Sir Cuartes N. E., K.C.M.G., C.B., F.Z.S.

Chromodorids from the Red Sea, collected and figured
loyy Mitre, Cyysenll Oreogsllenaelee (Vee weey ILI) ccoasgnenonconaeocone

FantuamM, Haroxtp B., D.Sc., B.A., F.Z.8., and Porrsr,
Miss Annie, D.Sc.

On a Bee-disease due to a Protozoal Parasite (Vosema

Harrison, Ruru, Oxford.

Some Madreporaria from the Persian Gulf. With
a Note on the Memoir and some Further Notes
on Pyrophyllia inflata by Sypnny J. Hickson, M.A.,
D.Se., F.R.S., F.Z.S8. (Plates LVII. & LVIII. and Text-
figs. 216 -221.)....:

Hetimayr, C. E., Division of Birds, Zoological Museum,
Munich.

A Contribution to the Ornithology of Western
Colombia

Sihohoneseueless}ens}aieei sie) elese/eleleieielolsieleleisidie tiles) s/s late elelelnlisielsfolulateletalsichalalefersle

Hickson, Sypvey J., M.A., D.Sc, F.R.S., FZS. See
Harrison, Rutu.

Some Madreporaria from the Persian Gulf,

Page

S07

1068

625

1018

1084

xili
Ho.pine, R. E.

Exhibition of the Horns of a Highland Ram, a Fallow
Deer, and a Roebuck, which were fused at the base,
and the skull of a coursing Greyhound with abnormal

COTA GEG T OTIS ee ee ec ee ae a eR Aer. Sy ROLE

HopeEweE.u-SmirH, A., L.R.C.P., M.R.C.S.,and Tims, H.. W.
Marett, M.A., M.D., F.L.S., F.Z.8.

Tooth-germs in the Wallaby (Macropus billardier?).
(Bla tee xeliValiian diiexct “mies SIE G99) ls eeeeee sees eon

LANKESTER, Sir E. Ray, K.C.B., F.R.S., F.Z.S.

Exhibition of a special supplement to the ‘ Field’ news-

PBAPOV ocr e reece cece cece eee eee ee eee eee t eee c ence entre eee seeetees

Lerrrer, Rosert T., D.Sc., M.B., F.Z.S.

A Demonstration of Nematode parasites obtained from
animals in the Zoological Gardems .....:.......-.0c0cc.ee-0

ListEer, JoserH J., M.A., F.R.S., F.L.S., F.Z.S.

The Distribution of the Avian Genus Megapodius in the
Jee rouire, Ikslenaglsyy f(UMeraratves N50.) Gs sccncacndandocaosecedeqoesse

LYDEKKER, RicHARD.

An African Rhinoceros, Klipspringer, and Gazelle.
(Mextatigss VOLSIOS jak. esac. doussais Maar ohana nt

Manpers, Lieut -Colonel Nevitie, R.A.M.C., E.Z.8., F.E.S.

An Investigation into the Validity of Miillerian and
other Forms of Mimicry, with Special Reference to the
Islands of Bourbon, Mauritius, and Ceylon. ...............

Page

946

926

620

749

958

X1V

Page
Meruvuen, The Hon. Paut A., F.Z.S.
On an Amphipod from the Transvaal. (Plates XLIX.—
Jill) ae at Aas A AAR 1A AM RE AE ESE SNR RB cs za 948
Mircuett, P. Cuatmers, M.A., D.Sc., Hon. LL.D., F.B.S.,
F.Z.5S., Secretary to the Society.
Report on the Additions to the Society’s Menagerie
ching auinemmontinvot sue bruce lO) Ibe ae ernest ene nee nene 557
A letter from the Governor of the Seychelles on Land-
Hortorsesminkthosewlislancds 1c eer ee eee eee et Renee 622
Report on the Additions to the Society’s Menagerie
diuiemsuthemmonthweteMarchenl 9/0 ii asses eeeet eee nein anne 671
Additional reference to the memoir “On Longevity
and Relative Viability in Mammals and Birds” ......... 868
Report on the Additions to the Society’s Menagerie
ChB HAS WS TOD OH Wyaall WOM jo yseassacacso9osa9cugcosoce 869
Report on the Additions to the Society’s Menagerie
GlpuetNS tae rmo ain Wey WAI 5.5, co2ccons00ascn000q0osoa000 985
Remarks upon the eggs of Struthio massaicus, S. aus-
tralis, and 8. molybdophanes, seen at Mr. Carl Hagenbeck’s
OStrichyHarmatastellancenyeeeeeek: cote ree eens ee enn eeeeener 987

Remarks upon a pair of young African Rhinoceros seen
at Mr. Carl Hagenbeck’s Zoological Park at Stellingen... 987
Morcnu, J. A., Christiania.

On the Natural History of Whalebone Whales. (Text-
Fes OOS O32) a) peissinddwacieg « semsiss cd sruscacte ce ache eet eee eee 661

Moss, The Rev. A. Miuus, M.A., F.Z.S., F.E.S.

Notice of a memoir on the “Sphingidee of Peru” ...... 869

XV
Niconi, WinuiaM, M.A., D.Sec., M.B., F.Z.S8.

On a unique Pathological Condition in a Hare. (Text-
HO LOD. Ay etme te Nees tds ce eorrk esta ce wed aw ig tale octane
On Three New Trematodes from Reptiles. (Plates
XXVIII: & XXVIII.)

Primmer, Henry G., F.R.S., F.LS., F.Z.8., Pathologist to
the Society.

Report on the Pathological Examination of Rats (J/us
decumanus) caught in the Regent’s Park and in the
Society’s Gardens

CEDOOIOIOIOIO POI IOC On ICICI ICO CICICICICECICEC IAI ICICNCECECECICICECHOIC

Pocock, Reernaup I., F.RS., F.LS., F.Z.8., Curator of
Mammals and Resident Superintendent of the
Gardens.

Exhibition of the skin of a new Chacma Baboon (Papio
porcarius, Subsp. griseipes), a specimen of the North-
American Black-footed Polecat (Putorius nigripes), ete.
eT MA Pi oiec aie oes occa intone sc on em nate seemnbenanos tthe

Exhibition of a newly born Masked Palm-Civet
(Paradoxurus larvatus), with an abnormal leg. (Text-

fig. 147.)

i i i ii i iii i i ii i ki i i i iii ii i i i ii i rice i i er ire aay

Exhibition of some hair from the “ puppy-coat” of a
Grey Seal (Halicherus grypus)

On the Palatability of some British Insects, with Notes
on the Significance of Mimetic Resemblances. With
Notes upon the Experiments by Prof. E. B. Potxron,
TS es Re ASie ea th oS ne aics dioltaiate ok EMU AN ar ci MR NE TaN

Exhibition of, and remarks upon, the skin and skull of
a specimen of the Crested Rat (Lophiomys ibeanus Thos.).
(Text-fig. 190.)

Oe ee ee ee i i i ee a

Page

674

677

558

621

696

809

XV1
Pocock, Reema I., F.RS. &e. (Continued.)

Exhibition of photographs of, and remarks upon, two
hybrid foals, Hquus asinus somaliensis x Equus zebra
and Hguwus asinus somaliensis x Equus quagga chap-
manm. (Text-figs. 201-203.)

Porter, Miss Anniz, D.Sc., and Fanrnam, Haron B.,
D.Sc., B.A., F.Z.S.

‘On a Bee-disease due to a Protozoal Parasite (Nosema

Poutton, Epwarp B., M.A., F.B.S., F.Z.8. See Pocock,
R. B.

On the Palatability of some British Insects, &e.

Reoay, C. Tare, M.A., F.Z.8.

Exhibition of a series of lantern-slides of scales of the
SHUMMTOM ((SOUMMO STGP) ococenrncesoepakocouacoocacvs.5hove0s06000 000

Seru-Smitn, Davin, F.Z.8., Curator of Birds.

Exhibition of a living Hybrid between the White-
eyed Pochard (Aythya nyroca) and the Marbled Duck
(Marmaronetia angustirostris) ............00cceceeseesseess eee

Exhibition of a nest of the Grey Struthidea or Apostle-
Bird (Struthidea cinerea), and lantern-slides of Penguins
in moult and of wild Swainson’s Lorikeets (7richoglossus
novee-hollandic). (Text-fig. 164.) ............cccceceseeeenes

Exhibition of, and remarks upon, two immature
Black-backed Porphyrios (Porphyrio melanonotus) with
wine=claws: (ext tis. 2005) vs. cceanss sis easeeen ce doemnCn eer

Page

988

625

671

5d8

671

XV
Sexron, Mrs. KE. W., Marine Biological Laboratory, Ply-
mouth.

On the Amphipod Genus Leptocheirus. (Plates XVII.—
SXGIDXE: erm cleaner ule Oa) yee time ae geen oh aaron Maries

SHUFELDT, Ropert W., M.D., C.M.Z.S.

Exhibition of a photograph of a male albino Woodchuck
(CAP GHONDUIS TOOINRD) “pa oencobe neater MneuGsoncdoe echoes eh coHCBr enn

SMALLEY, FrepERIc W., F.Z.S., and Bonuorr, J. Lewis,
Wi oBloe TIGIUES 5 1WZAdse

On Colour and Colour-pattern Inheritance in Pigeons.

(Tete n gets 6) GD EPONA Ei os Ahen Nn ORE coc nce ese meee
Suirn, A. Hoprweni-. See Hopmweiu-Surru, A.
Smit, D. Seru-. See Sera-Surru, D.

THomas, OLDFIELD, F.R.S., F.Z.8.

The Duke of Bedford’s Zoological Exploration of
Eastern <Asia.—XIV. On Mammals from Southern
Shen-si, Central ‘China. ‘(Plate X XTX.) 1....0.0.0.....4..

Tomson, J. Sruart, Ph.D., F.R.S.E., F.L.S., Lecturer
and Senior Demonstrator in Zoology, University of
Manchester.

The Alcyonaria of the Cape of Good Hope and Natal.—
GorconacEa, (Plates XLITI.—XLV. and Text-fig. 167.)
Tims, H. W. Marerr, M.A., M.D., F.L.S., F.Z.S:, and
Horrwett-Suiru, A., L.R.C.P., M.R.C.S.
Tooth-germs in the Wallaby (Macropus billardieri).
(Plens CIEL. eiavel AMexceileiss JUS IS) s) gcse onceckoqubapacod

Proc. Zoou. Soc.—1911., 5

946

60]

XVI

NEW GENERIC TERMS

PROPOSED IN THE PRESENT VOLUME (pp. 557-1218).

Page Page
Anoplotenia (Vermidea) ...... 1003 Eriphostoma (Reptilia)......... 1078
Arctognathns (Reptilia) ...... 1079 Ictidognathus (Reptilia) ...... 1078
Arctosuchus (Reptilia) ......... 1079 Moschops (Reptilia) ............ 1073
Dasymetra (Vermidea) ......... 683 Taognathus (Reptilia) ......... 1076

Dizelurodon (Reptilia) ......... 1075 Thysanoteenia (Vermidea)...... 1002

SCLENTIFIC

Abraxas, 867.
grossulariata, 833.
Acanthogorgia
armata, 870, 880.
Acanthopneuste
viridanus, 718.
Accipiter
superciliosus, 1203.
tinus, 1203.
Aceratherium, 945.
Acontias, 705.
burtoni, 705.
Acrea, 714, 742,
Acridotheres
cristatellus, 860.
tristis, 698, 701, 720,
740.
Acrocephalus
dumetorum, 718.
stentoreus, 718.
Acrorchilus

erythrops griseigularis,
1149.

Acryllium
vulturinwn, 862.

Actinote, 705.

Adelomyia
cervina, 1211.

melanogenys maculata,

1211.
AXgithina
tiphia, 716.
A®lurosaurus, 1079.
fetinus, 1077, 1081.
tenuirostris, 1077,
1081, 1082.
whaitst, 1077, 1078,
1081, 1082.
fflurosuchus, 923,
924.
/#lurus
fulgens, 557.
Aapyprymnus, 940,

INDEX

OF

Acelecyathus
persicus, 1019, 1034,
1035, 1039.
Aglaia
Fanny, 1107.
gyroloides, 1104.
peruviana, 1104.
Agyrtria
francie, 1182.
Alcadia, 761, 763, 776,
777, 780, 782, 783,
785, 794, 796, 797,
799, 800.

hollandi, 762, 766, 770,
TOE, TUB, HE WAY
781, 785, 787, 788,
793, 798, 807, 808,

809.

palliata, 762, 766, 768,
769, 770, 772, 773,
778, 787, 788, 789,

793, 798, 807.

Alcedo

amazona, 1192.

americana, 1192.

cabanisti, 1198.

inda. 1192.

torquata, 1192.
Algiroides

nigro-punctatus, 864.
Allantus

arenatus, 828, 850.
Aloposaurus, 919,

1078.

gracilis, 1081.
Alseonax

latirostris, 720.

muttut, 720.
Alytes

obstetricans, 696.
Amazilia

Juscicaudata, 1183.

riefferi, 1188.

NAMES.

Amazilia
tzacatl fuscicaudata,
1183.
— yucunda,
1183
Amblycercus
holosericeus, 1122.
Ampelis
natterertt, 1146.
Anabates
subalaris, 1151.
Anabazenops
lineatus, 1151.
mentalis, 1151.
subularis, 1151.
Anas
cyanoptera, 1209.
superciliosa, 755.
Androdon
equatorialis, 1176,
1177.
Anonchoteenia, 637.
Anoplocephala, 651,
653, 660.
Anoplops
bicolor, 1171.
— equatorialis, 1171,
1172, 1211.
— bicolor, 1171, 1173,
1212.
— dague, 1170, 1171,
1173, W211, 1212.

1182,

— olivascens, 1171,
1173.

— ruficeps, 1171,
1173.

leucaspis, 1171,
1172.

Anoplotznia, gen. nov.,
1003, 1018.

dasyuri, 1003-1007,

1009, 1010, 1012,
1013, 1OL5.

82 *

xx

Anthrocera,
(Zygena) filipendule,
832.

Anthus
rufulus, 722.
Antilope
corrina, 962.
Aphanoconia, 761, 780,
785, 796, 797, 798.
andamnanica, 762, 773,
787, 801, 809.
gouldiana, 762, .7685,
778, 779, 785, 793,
808, 809.
merguiensis, 762, 779,
501, 808, 809.
pachystoma ponsonbyi,
763

rogersii, 762, 793, 801,
809.

(Helicina) theobaldiana,
60.

Aphantopus
hyperanthus, 8217,
828.
Aphelocoma
woodhousei, 869.
Apis

mellifica, 812, 850.
Aploparaksis, 652.
Apocrangonyx, 949,
Apodemus

aqrarius pallidior, 690.

speciosus peninsula,

690.
Appias, 730, 734, 787,
742.

galene, 709, 735.

paulina, 730.
Aptenodytes

pennant?, 558, 671.
Aramides

mangle, 1209.

wolfi, 1208, 1209.
Araschnia

levana, 823, 865.

prorsa, 823, 865.
Arctognathus, gen. nov.,

1079.

curvimola, 1079.
Arctomys

monax, 946.
Arctonyx

leucolemus, 688, 689.

— orestes, 688.
Arctophila, 854.

mussitans, 848, 851,

853.
Arctosuchus, gen. nov.,
1079.
tigrinus, 1079.

Argiades
sylvanus, 831.
Argilleecia
affinis, 595, 596,
601.
Argynnis
aglaia, 824.
castetst, 725.
euphrosyne, 820, 826.
hyperbius, 702, 703,
704, 706, 709, 735,
736, 737.
(Brenthis) ewphrosyne,
824,
(—) selene, 825.
(Dryas) paphia, $24,
827, 828.
Arion, 804.
ater, 813
hortensis, 813.
Arremon
aurantiirostris, 1118.
— erythrorhynchus,
1119.
— occidentalis, 1118,

— spectahiiis,
1119.

erythrorhynchus, 1118,
1119.

1118,

occidentalis, 1119.
spectabilis, 1118, 1119.
Arrenga
blight, 716.
Artamus
Fuscus, 720, 729.
personatus, 859.
superciliosus, 859.
Artemia
salina, 986.
Ascaris
canis, 620.
Aspidosiphon, 1023,
1025, 1027.
Aspidura sp., 716.
Asturina
schistacea, 1204.
Aulacorhamphus
albivitta, 1213.
— pheolemus, 1218.
ceruleiqularis,
1218.
petax, 1213.
pheolemus, 1213.
sulcatus, 862.
Aurelia
aurita,
1056.
Automolus
assimilis, 1150.
holostictus, 1152.

1047, 1053,

INDEX OF SCIENTIFIC NAMES.

Automolus

nigricauda, 1150, 1210.

(Rhopoctites) 7fo-

brunneus, 1149, 1150.

Avicula

zebra, 1058, 1059, 1060.
Aythya

nyroca, 5d8.

Bacillus
pestiformis, 626.
Bairdia
acanthigera, 595.
amygdaloides, 595.
dubia, 595, 596, 601.
mediterranea, 595.
obtusata, 595.
Balena
australis, 663.
Baleenoptera
borealis, 662, 668.
musculus, 662,
sihbaldii, 662.

Balanus
improvisus, 591.
Balistes
flavimarginatus, 1058,
1065.
viridescens, 1058, 1065
Basileuterus
auricularis, 1092.
Julvicauda semicer-

vinus, 1092.
melanotis, 1092.
— dedalus, 1092.
meridanus, 1092.
semicervinus, 1092.
tristriatus, 1092.
— melanotis, 1092,
1212.
— meridanus, 1092.
— tristriatus, 1092,
1212.

Bauria, 898, 899, 900,
901, 902, 905, 909,
913, 914, 915, 916,
917, 920, 923.

cynops, 895, 896, 897,
924, 925.
Belone sp., 1064.
Bertia
satyri, 649.
studeri, 649.
Bertiella, 994, 999, 1004,
1017.
cercopithect, 638, 640,
641, 643, 644, 647,
649, 650, 651.
conferta, 645, 646, 650,
edulis, 1017.

Bertiella
melanocephala, 639,
640.
mucronata, 639, 645,
646, 650. -

obesa, 1017.
plastica, 650.
polyorchis, 646, 651.
sarasinorum, LOL.
semonti, 1017.
Betta
splendens, 989.
Bison
bonasus, 557.
Bistylia, 1080.
Boeckia
typica, 563.
Boisscunneaua
flavescens flavescens,
1185.
— tinochlora, 1185.
Bombus, 841, 853.
agrorum, 846, 850,
803

hortorum, 812, 852,
854, 855.
joncellus, 891.
lapidarius, 845, 880,
852.
® terrestris, 851, 854.
Boreophausia
ineriis, 666, 667.
Bothriocephalus, 639.
Brenthis, 865.
Buarremon
brunneinucha, 1117.
Bubo
poénsis, 869.
Bucco
macrodactylus, 1196.
noaname, 1195, 1196,
1211.
pectoralis, 1194, 1195,
Uzi

subtectus, 1195.

tectus, 1195.

— subtectus, 1195,

We,

— tectus, 1195.
Buchanga

atra, 7217.
Bucorax

abyssinicus, 862.

caffer, 862.
Budorcas

bedfordi, 693, 695.

mitchelli, 695,

INDEX OF SCIENTIFIC NAMES.

Buthraupis
arcet, 1110, 1212.
aureocincta, 1110,
T1111, 1211.

edwardsi, 1111, 1211.

melanochlamys, 110,
1110, 1211, 1212.

rothschildi, 1110, 1211.

Bythocypris
reniformis, 595.

Cabera
exanthemaria, 832.
pusaria, 833.
Cabreta
leschenaultii, 706.
Cacicus
microrhynchus, 1212.

uropygtatis, 1121, 1212.

Cacomantis
passerinus, 723.
Calanus, 668.
Calliphora
vomitoria, 850.
Calliste
aurulenta, 1102.
cayanda, 860. .
emilie, 1103.
Jfrancesce, 1107.
Julvicervia, 1109.

gyroloides, 1104, 1107.

icterocephala, 1198.

Johanne, 1101.

labradorides, 1109.

larvata, 1108.

lavinia, 1103.

leucotis, 1109.

ruficervix, 1109.

rufigularis, 1102.

taylori, 1109.
Callithrix

Jacchus, 856.
Calophasis

ellioti, 862.
Calospiza

aurulenta aurulenta,

1102.

— sclateri, 11C3.

florida arcet, 1212.

— florida, 1212.

gyroloides, 1105.

— bangsi, 1105, 1106,

1107, 1212.
— catharine, 1106.
— deleticia, 1104.
— gyroloides, 1104,
1105, 1106, 1212.

XX1

Calospiza

larvata fanny, 1107,
1108, 1212.

— francesce, 1108.

— larvata, 1108.

lavinia, 1103, 1104.

—dalmasi, 1104,
1212.

— lavinia. 1103, 1105,
1212.

palmeri, 1107, 1211,

phenicotis, 1211.,

ruficervix, 1108.

— fulvicervix, 1109.

— ruficerviz, 1109.

— taylort, 1109.

rufigula, 1102.

Calotes, 727.

nigrilabris, 708.
ophiomachus, 707, 708,
739

polytes, 707.
versicolor, TOT, 708,
Ws 7

Calyptrophora

josephine, 886.

Canielus

bactrianus, 869.

Campophaga

sykesi, 719.

Capito

equatorialis, 1200.

bourcierti, 1199.

— eguatorialis, 1200.

— bourcierti, 1199,
1200.

— salvini, 1199, 1200.

maculicoronatus, 1198,
1199, 1211.

quinticolor, 1198, 1211.

shelleyi, 1200.

sguamatus, 1199, 1211,

Capra

cylindricornis, 973.
hispanica, 963, 970,

ibex, 963.

pyrenaica, 963, 964,
967, 969, 970, 972,
974, 975, 976.

— hispanica, 963, 966,
967, 969, 970, 972,
975, 976, 977.

— pyrenaica, 969, 970,
975.

— victorie, 975, 977

stbirica, 969.

sinensis, 695.

taxicolor, 694.

tibetanus, 687, 693,
694, 699. .

icterocephala, 1103.

johanne, 1101, 1102,
-1210, 1212.

labradorides, 1109.

Capreolus
bedtordi, 695.

Caprimulgus
rosenberg?, 1210,

Xx11

Carabus, 842.
violaceus, 816, 836,
841, 845.
Caranx sp., 1069.
Carbo
brasilianus, 1210.
vigua, 1209.
Cariama
cristata, 864.
Carpodectes
hopkei, 1147, 1210,
1212, 1218.
nitidus, 1212.
Caryophyllia
communis, 1088.
Casella
atromarginata, 1062.
Cassiculus
microrhynchus, 1121,
22.
Cassicus
persicus, 860.
pyrohypogaster, 4123.

uropygialis, 1121,1122. |

Cassidix

oryzivora, 1122.

—- oryzivora, 1122.

— violea, 1122.
Catenotenia, 637, 1018.
Catheturus

lathami, 863.
Catopsilia, 730, 734, 742.

crocale, 725.

florella, 727.

pomona, 709.

pyranthe, 727, 738,

739, 741.
Cebus sp., 815, 825, 829,
831, 832, 851, 856.

capucinus, 651.

loricatus, 1188, 1189.

- mentalis, 1189.

squamatus, 1189.
Ceophlcus

lineatus lineatus, 1189.

Cephalophus
ignifer, 673.
johnstont, 672.
weynst, 672, 673.

Ceratodus, 988.
forsteri, 942.

Ceratogymna
atrata, 861.
elata, 861.

Ceratoisis
ramosa, 870, 877, 892.
Ceratophora

myslaceus, 706.
negrilabris, 706, 708.
ophiomachus, 706.
steddarti, 708, 710.

Ceratophora
versicolor, 706.
Ceratopoma, 761.
Ceratosoma
cornigerum, 1068, 1065.
Cerchneis
punctata, 700.
Cercomacra
berlepschi, 1167,
1210.
carbonaria, 1166.
crepera, 1165.
maculicaudis, 1166.
nigricans, 1166.
tyrannina, 1165, 1167.

— rifiventris, 1165,
1166.
— tyrannina, 1169.
Cercopithecus

callitrichus, 688, 649.

centralis, 671.

denti, 856.

mona, 856.

pygerythrus, 856.

roloway, 856.
Certhia

cerulea, 1097.
Certhiola

chloropygia, 1098.

mexicana, 1098.
Cervus

bedfordi, 560.

hortulorum, 558.

canthopygus, 560.
Ceryle

amazona, 1192.

americana americana,

1192, 1198.

— cabanisii, 1193.

cabanistii, 1192.

inda, 1192.

torguata, 1192.

— torquata, 1192.
Cethosia

nietnert, 703.
Cheetura

indica, 7238.
Chaleidoseps, 705.

thwaitesir, 706.
Chalybura

buffont, 1184, 1185.

tsaure, 1212.

urochrysa, 1184, 1185.

1212.

Chama, 1059.

sp., 1057, 1061.

Joliata, 1057, 1061.
Chameleon, 922.

sp., LOG6.
Chamepelia

minuta, 869.

INDEX OF SCIENTIFIC NAMES.

Chameepetes
goudoti, 1207.

— goudotii, 1207.

— rufiventris. 1207,
Chapmannia, 652.
Charaxes

imna, 726.

Chelone, 921.
imbricata, 624.
mydas, 624.

Chilosia
illustrata, 848, 853,

854, 855.

Chiromacheris

aurantiaca,
1212.

vitellina, 1141, 1211,
PAN

Chlamydodera
naculata, 861.

1141.

| Chlorochrvsa

nitidissima, 1101,
1211.
phenicotis, 1211.
Chloronerpes
cecilie, 1188.
gularis, 1187.
rubiginosus, 1187.
— gularis, 1187, 1211,
1212.
— rubiginosus, 1187.
— uropygialis, 1212.
Chlorephanes
atricapilla, 1097.
cerulescens, 1097.
spiza . cerulescens,
1097.
— exsul, 1096, 1097.

| Chloropipo

holochlora, 1138.
— lite, 1138, 1210.

| Chloropsis

jerdoni, 716.
malabarica, 716.

Chlorostilbon

angustipennis, 1183.
pumilus, 1188.
Chlorothraupis
olivacea, 1114.
Chromodoris sp., 1070.
annulata, 1063.
diardui, 1062,
1064.
elizabcthina, 1062.
hilaris, 1063.
inopinata, 1070, 1072.
nigrostriata, 1062
quadricolor,1062.
reticulata, 1063, 1064,
1065, 1068, 1069,
1070, 1072.

1063,

Chromodoris

sykest, 1063.

tinctoria, 1069, 1072.
Chrysomela

polita, 841.
Chrysophanus

phleas, 830.
Chrysophrys

bifasciata, 1065.
Cicinnurus

regius, 861.
Cistecephalus, 1080,

1081.

microrhinus, 1081.
Cisticola

cursitans, 718.
Cistudo

carolina, 624.
Cittocincla

macrura, 721, 857.
Cladius

viminalis, 849.
Clavatella

(Eleutheria) prolifera,

1053.

Clematissa, 880.
Climacocercus
zonothorax, 1203.
Clisiocampa
neustria, &32.
Clytia, 1046.
folleata, 1048.

Cnipodectes

minor, 1128, 1129,
1130.
subbrunneus, 1128,

1129, 1180, 1213. »
— minor, 1129, 1130.
— subbrunneus, 1128,
1129.
Cobus
cob, 674.
Coccidium
cuniculi, 676.
Coccinella, 816, 845.
7-punctata, 812, 819,
841, 842, 845, 847.
Coccystes
coromandus, 724.
jacobinus, 724.
Coccyzus
melacoryphus, 1202.
Ceenonympha
pamphilus, 825, 829.
Ceereba
cerulea, 1097.
— luteola, 1098.
— microrhyncha, 1097,
1098.
chloropyga mexicana,
1098. ;

INDEX OF SCIENTIFIC NAMES.

h

Colias
nilgiriensis, 72d.

— Collocalia

francica, 698, 699.

' Colluricinela

heinet, 754.
Collyriocinela
harmonica, 859.
Columba
albilinea, 1205.
— albilinea, 1205.
— crissalis, 1205.
albitineata, 1205.
goodsom, 1205, 1206,
1210, 1218.
nigrirostris brunnet-
cauda, 1205, 1213.
plumbea bogotensis,
1206.
speciosa, 1205.
subvinacea berlepschi,
1206.
Compsocoma
notabilis, 1111.
Conchoderma
auritum, 667, 668.
Conocyathus, 1030.
zelandie, 1019, 1020,
1029, 1032, 1044.
Conopophaga
castaneiceps, 1176.
brunneinucha,
1176.
—- castaneiceps, 1176.
gutturalis, 1176.
nevutoides, 1167.
Conosmilia, 1088,
1041.
anomala, 1039.
elegans, 1039.
granulata, 1040,
lituolus, 1039.
striata, 1040.
stylifera, 1040.
Cophotis
zeytanica, 706, 708,
710.
Copsychus
saularis,
857.
Copurus
leuconotus, 112d.
Coracias
indica, 722.
Cordulegaster
annulatus, 835.
Coronula
diadema, 667, 668.

1039,

Ts TBR

| Corvus

cornia, 861.
scapulatus, 861.

XXU1

Coryphotriccus
albovittatus, 1134.
— distinctus, 1135.
parvus, 1139.

Cosmotricha
potatoria, 834.

Cossus
ligniperda, 831.

Cossypha
caffra, 857.

Cotinga,
cayana, 1147.
natterertt, 1146, 1147,

1211.
ridgway?, 1146, 1147.
simont, 1146.

Cotyle
uropygialis, 1093.

Cracticus
destructor, 861.

Crangonyx, 949, 951.
flageliula, 950.
subterraneus, 957.

Craspedoprion
equinoctialis, 1127.
brevirostris, 1127.
olivaceus, 1127.

Crateropus
canorus, 716.
cinereifrons, 715.
rufescens, 715.
striatus, 715.

Cratopus, 701.

Crax
alector, 863.
caruncuiata, 863.
globicera, 863.

Creciscus
alhigularis, 1208.

Cricetomys
gambianus, 651, 9995,

1003.

Cricetulus
andersont, 691.
triton, 691.

Crithidia, 626.

Crocidura
attenuata, 688.
coree, 688.

Crossarchus
fasciatus, 856.

Crosslandia
viridis, 1064.

Cuculus
canorus, 723.
micropterus, 723.
poliocephalus, 723.

Culcitella, 652.

Culicicapa
ceylonensis, 721.

| Cuscus, 921.

XX1V

Cyanerpes
cerulea microrhyncha,
1097.
Cyanocitta
pulchra, 1124.
Cyanocorax
affinis, 1128, 1124.
— affinis, 1123, 1124,
1212.
— zeledoni, 1124, 1212.
pileatus, 1124.
sclateri, 1124.
Cranolesbia
kingit, 1187, 1211.
— celestis, 1187, 1211.
— emme, 1187.
— kingit, 1187.
Cyanolyca
armillata, 1124.
pulchra, 1124.
Cyanomyia
francie, 1182.
Cyclorchida, 652.
Cyclorhynchus
e@quinoctialis, 1127.
cinereiceps, 1126.
subbrunneus, 1128,
1129.
Cycloseris
hexagonalis, 1036.
Cymbilanius
lineatus, 1157.
— fasciatus, 1157,
Cynanthus
celestis, 1187.
Cynochampsa
laniaria, 893.
Cynodraco
serridens, 893.
Cynognathus, 895, 899,
900, 904, 906, 907,
908, 910, 912, 923,
925, 1080.

berryi, 902, 905, 925.

erateronotus, 894, 900,
905.
leptorhinus, 901.
platuceps, 894, 901,
903.
seeley?, 893, 902.
Cyornis

rubeculoides, 720.

tickelli, 720,
Cyphorhinus

brunnescens, 1088
Cyphorinus

cantans, 1089.

lawrencii, 1039.

pheocephalus, 1088.
Cypselus

affinis, 723.

Cypselus
melba, 723.
Cythere
albo-maculata, 595.
cingulata, 595, 596,
601.
convexra, a95.
crispata, 595, 596, 601.
diffusa, 596.
elegans, 596.
emaciata, 595.
tuberculata, 595.
Cythereis
antiquata, 595.
deformis, 5965,
601.
Jonesit, 595.
runcinata, 595.

597,

Cytherella

ovalis, 595, 600, 601.
Cytheridea

elongata, 595.
Cytherideis

subulata — crenulata,

595, 600, 601.
— fasciata, 600.
Cytherura
cellulosa, 595.
eribriformis, 595.

cribrosa, 595, 599,
601.

fossulata, 595, 599,
Ol.

maculosa, 595, 599,
601.

striata, 595.

Dacelo
cervind, 861.
gegantea, 861.
leachii, 861.

| Daenis

cayana, 1095, 1096.

— callaina, 1096.

— cayana, 1096.

— cerebicolor,
1096, 1212.

— glaucogularis, 1096.

— napeéa, 1095, 1096.

— ultramarina, 1096,
1212,

cerebicolor, 1095, 1096.

napea, 1096.

salmoni, 1116, 1117.

venusta, 1094.
fiiginata,

1212.

Danais, 742.
aglea, 702, 725.
ceylonica, T7038,

1095,

1094,

INDEX OF SCIENTIFIC NAMES.

Danais
chrysippus, 702, 703,
704, 708, 725, 727,
731, 738, 739, 740,
744, 745.
Sumata, 709, 735, 736.
limniace, 704, 725, 737.

plexippus, 702, 703,
704,

septentrionis, 704, 725,
726, 727

( Amaur is) eee 702.
Dania

rero, 985.
Dasymetra,

683, 685.

conferta, 683, 686.
Dasy pus, 919.

villosus, 857.
Dasyurus

ursinus, 1003, 1017.
Davainea, 639, 652.
Dawsoniella, 762

gen. noyv.,

Delias 4
eucharis, 703, 708, 737,
740.
Delphinognathus, 919,

1074, 1075
conocephalus, 1073.
Dendrochelidon
coronata, 727.
Dendrocinela
atrirostris, 1156.
erythropygia, 1155.
Jumigata, 1156.
lafresnayet, 1155, 1156.
— pheéochroa, 1155.
meruloides, 1155.
— aphanta, 1156.
— lafresnayei, 1155,

1156.
— meruloides, 1155.
— pheochroa, 1155,
1156.

olivacea, 1155, 1156. .

— anguina, 1157.

— lafresnayei, 1155,

1156.

— pheochroa, 1156,

pheochroa, 1156.
Dendrocolaptes

atrirostris, 1156.

cuneatus, 1152.

triangularis, 1153.
Dendrocops

meruloides, 1155.
Dendrocopus

major, S61.
Dendrocygna

autumnalis, 1209.

discolor, 1209.

INDEX OF SCIENTIFIC NAMES.

Dendroica

estiva, 1091.

— estiva, 1091.

castanea, 1091.
Dendrohyrax, 673.
Dendrophyllia sp., 1019,

1036, 1039, 1044.
Dendropupa, 803.
Dendrornis sp., 1153.

equatorialis, 1153.

erythropygia @equa-

torialis, 1158.
lachrymosa, 1153.
— lachrymosa, 1212.
— rostrata, 1153, 1210,
1212.

punctiqula, 1153.

triangularis — e@qua-
torialis, 1153, 1154,
1212.

— punctigula, 1154,

— triangularis, 1154,
1155.

Dendrortyx

leucophrys, 863.
Dermochelys, 921.

coriacea, 9205.
Deuterosaurus, 1075.
Diadectes, 920.
Diademodon, 894, 923,

924, 920.

mastacus, 908.

Dizlurodon, gen.

1075.
whaitsi,
1082.
Diallactes

granadensis, 1158..
Dicheilonema

horrida, 620.

Dicrurus, 725.

ater, 717.

leucopygialis, 718, 728.

longicaudatus, 717.
Dictyocaulis

filaria, 620.

Dicynodon, 893, 1075,
1080, 1081.

sp., LOSI.

Didelphys, 921, 941,1017.
Diglossa

cerulescens, 1093.
Diglossopis

cerulescens, 1093.

— cerulescens,

_ 1094.

— pallida, 1094.
Dilepis, 652.
Dinotherium

curviceri, 944.

nov.,

1075, 1081,

1093,

Dinotherium
giganteum, 944.
hobleyi, 943, 944, 945.

Diorchis, 652.

Dissemuroides
lophorhinus, 718.
paradiseus, 718.

Disythamnus
rufiventris, 1165.

Dorcelaphus
savannarum, 869.

Doris
tinctoria, 1070.

Dromatherium, 924.

Dryocopus
lineatus, 1189.

Dryotriorchis
spectabilis, 869.

Dumetia
albigularis, 715.

Duneania, 1041.

Eccasaurus, 1075.
Echidna, 918, 919, 921.
aculeata typica, 924.
Echinomyia
ferox, 848. 855.
Elenia
cayenennsis, 1134.
cinerea, 1132, 11388.
— cinerea, 1132.
— parambe,
1183, 1210.
Elaphrornis
pallisert, 716.
Elymnias
fraterna, 702.
Embernagra
brunneinucha, 1117.
Empidonax
acadicus, 1186.
virescens, 1186.
Em pis
tesscllata, 848.
Enys
sculpta, 624.
Endothiodon, 1080.
bathystoma, 1081.
wniseries, 1075, 1081.
Engraulis
boelama, 1064.
Entameeba
apis, 626.
coli, 626.
Epenthis, 1048.
Hipimys
confiucianus, 689, 690.
— canorus, 690.
— luticolor, 689, 690.
— sacer, 690..
luticolor, 690. . »

1132,

XXV

Epimys
sacer, 690.
Epinephele
Jartina, 816, 820, 828.
Janira, 828.
tithonus, 829.
Equus
asinus, 993.
— somaliensis, 98,
993.
grevyt, 998.
hemionus hemionus,993.
kiang, 993.
quagga, 993.
= Sai 993.
— chapmanni, 992.
zebra, 989, 993.
Eranna
gucunda, 1182.
EKresia, 705.
Eriocnemis
aurelie, 1185, 1186. |
Eriphostoma, gen. nov.,
1078, 1079.
microdon, 1078, 1082.
Kristalis, $12.
Erythrura
trichroa, 985.
EHucephala
humboldti, 1210.
Euchelia, 867.
jacobee, 815, 820, 825,
8382, 845, 845.
Euchloé °
cardamines, 815, 820,
865.
Eucometis
cassini, 1116.
Eucope, 1046,
1055, 1056.
Kucrangonyx, 948.
robertsi, 950, 951, 957.
vejdovskyt, 949, 950,
951.
Eucythere
prava, 595.
Eulipoa, 751.
Kunicella
papillosa,
881, 892.
Euphonia
orevirostris, 1101.
fulvicrissa, 1100.
— fulvicrissa, 1211.
— purpurascens, 1100,

1048,

871, 880,

PAIN
purpurascens, 1100.
avanthogaster, 1100,

1101

— brevirostris, 1101.
— brunneifrons, 1101.

XXV1

Euphonia
xanthogaster chocoensis,
1100, 1211.
— wanthogaster, 1211.
Euplexaura
albida, 884. *
brauer?, 884.
media, 871, 883, 8938.
parciclados, 884.
Euplea, 713, 729, 734,
739, 742.
core, 702, 708, 704,
708, 709, 720, 725,
726, 730, 737, 738
740, 741, 743, 744.
coreta, 702, 704, 725,
726.
euphon, 702.
goudoti, 698.
klugit, 702.
Hupodotis
australis, 868.
Euripus
consimilis, 703.
Kurypyga
helias, 864.
Euscarthmus
cinereus, 1180.
Eutoxeres
aquila, 1180.
— aquila, 1180.
— heterura, 1180,1181.
baront, 1180, 1181.
heterura, 1180.
salvint, 1180, 1181.
Eutrochatella, 761, 777,
778, 779, 780, 782,
783, 785, 788, 796,
798, 799.
pulchella, 762, 763,
768, 772, 773, 786,
787, 788, 793, 797,
807, 808, 809.

Falco
cachinnans, 1204.
cayennensis, 1205,
palliatus, 1204.
superciliosus, 12038.
Felis
concolor, 671.
Sontanieri, 688.
lyne isabellinus, 557.
microtis, 688,
serval, 869.

Filaria
australis, 620.
Flabellum

laciniatum, 1088.
magnificum, 1019,

1020, 1039, 1044.

INDEX OF SCIENTIFIC NAMES.

Flabellum

rubrum, 1021.
Florisuga

mellivora, 1181.

— mellivora, 1181.

Formica

rufa, 812, 849, 868.
Formicarius

anatis, 1173.

— destructus, 1173,

1174, 1212.
— nigricapillus, 1174,
1212.
destructus, 1174.
migricapillus, 1174.
rufipectus, 11'74.,
— rufipectus, 1174,
1175.

— thoracicus, 1175.
saturatus, 1174.
thoracicus, 1175.
Formicivora
consobrina, 1168.
melena, 1162.
quadrivittata, 1160.
quixensis —_ houcardi,
1164, 1212.
— consobrina,
1164, 1212.
schisticolor, 1163.
Foudia
erythrocephala, 701.
Franklinia
gracilis, 718.
Fregilupus
capensis, 699.
varius, 699.
Frinyvilla
rubra, 1118,
Fulica
martinica, 1208.
Fundulus
gularis, 985.
Fungia, 1042.
patella, 1019, 10386,
1089.

1163,

Galbula
melanogenia, 1194.
ruficauda, 1194.

Galeoscoptes
carolinensis, 857.

Galepus, 919.

Galesaurus,
(00

894, 899,
planiceps, 893, 898,
928, 925.
Galictis
vittata, 857.
Gallinula
mangle, 1209,

Gammarus, 949.

Garrulax
leucolophus, 859.
picticollis, 859.

Gazella
cuvieri, 961.
dorcas, 961, 962.
hayi, 961, 962.
kavella, 962.
leptoceros, 961.

Genneus
nycthemerus, 862.
Geocichla

citrina, 857.
(Monticola) eyanus,
857.

( — ) sawatilis, 857.
Georissa, 760, 800.
Geotrupes

vernalis, 839,

eotrygon

bourcieri, 1206.

veraguensis, 1206.

— eachabiensis, 1206,

1210, 12138.
— cachaviensis, 1206.
— veraguensis, 1206,
ac
Glaucidium

perlatum, 862.
Glaucis

e@neus, 1178.

columbiana, 1178.

Fraseri, 1177.

hirsuta, 1178.

— enea, 1178.
Glyphorhynchus

castelnaudit, 1152.

cuneatus, 1152.

— castelnaudit, 1152.
Gomphodontia, 894.
Gomphognathus, 894,

920, 923, 925.

rannemeyert, 908, 909,

910, 911, 912.
minor, 908, 909, 910,
911, 912.

polyphagus, 909.
Gonatodes

kandianus, 705.
Gonionemus, 1046, 1055.

Gorgonia

albicans, 881.

capensis, 871, 884,
887.

Hlammeoa, 871, 877,
888.

Gorgonocephalus, 874.
Gracula
intermedia, 860.
religiosa, 860.

Graculipica
melanoptera, 860.
Grallaricula
cucullata, 1176.
Grallina
australis, 858.
Graucalus
macit, 719.
Grison
Sura, 857.
Gryporhynchus, 692.
Guttera
pucherani, 863.
Guynia, 1038,
1040, 1041.
Gymnopithys
bicolor dague, 1170.
leucaspis, 1172.
ruficeps, 1172, 1173.
Gymnorhina
leuconota, 861.

1039,

Halcyon
sacra, T54.
smyrnensis, 723.
Halicherus
grypus, 696.
Hapalemur
griseus, 856.
Haplochilus
chapert, 985.
elegans, 985.
panchaz, 985,
Haplophyliia,
1041.
Harpactus
fasciatus, 723.
Harpalus, 836, 839.
ruficornis, 838.
Heleodytes
albobrunneus,

1211.

1038,

1088,

— albobrunneus, 1212. |

— harterti,
1212.
harterti, 1088.
Heliangelus
exortis, 1186.
Helicina, 772, 780, 783,
797, 798, 799.
brasiliensis, 760, 786,
787, 792.
japonica, 760, 784.

1038,

kubaryi, 59, 760,
Cae

sagraina, "760, 786,
787, 792.

substriata convexa, 761.
titanica. 7d9, 768,
786, 806.

INDEX OF SCIENTIFIC NAMES.

| Helicina

(Sturanya) beryllina,
760

| Heliconius, 742.

Heliothrix
auritus, 1186.
harrott, 1186.
Heliotrypha
parzudakii, 1186,
Helix, 762.
Helogale
varia, 557.
Hemipus
picatus, 719.
Hemithraupis
chrysomelas,
1212.
ruficapilla, 1117.
salmom, 1116, 1117,
1212.

IULN7/,

Henicorhina *
inornata, 1090, 1210,
1212;

leucosticta, 1090.
— eucharis, 1090.
— prostheleuca, 1090,
1212.
prostheleuca, 1090.
Hercynella, 803.
Herpetotheres
cachinnans, 1204.
Hesperia
antonia, 866.
carthami, 866.

side, 866.
Heterocyathus, 1021,
1031.
equicostatus, 1019,

1022, 1024, 1025,
1026, 1032, 1039,
1044.
alternatus, 1024, 1026,
1044,
cochlea, 1025.
heterocostatus, 1019,
1026, 1039, 1044.
lamellosus, 1025.
oblongatus, 1026.

parasiticus, 1025,
1026.

philippinensis, 1025,
1026.

pulchellus, 1026.

sulcatus, 1025.

wood-masom, 1026.
Heterodon

platyrhinus, 677.
Heteropsammia, 1031,

1039.
aphrodes, 1019, 1636.
michelini, 1032,

XXVli

Heterospingus
rubrifrons, 1115,
1212.
avanthopygius, 1114,

1115, 1212.
Hicksonella, 891.

spiralis, 871, 839.
Hirundo

chalybea, 1093.

ruficollis, 1093.

rustica, 722, 730.

viridis, 1099.
Hyalimax, 804.
Hyalonema, 877.
Hydrocena, 783,

800.

cattaroensis, 739, 760,

777, 806.
Hydrocorax
vigua, 1209.
Hyla

goughi, 1082, 1083.

misera, 1088.

rubra, 1032.

strigilata, 1083.

venulosa, 1082.
Hylocichla

mustelina, 857.

ustulata swainsonit,

1087.
Hyloctistes
subulatus — assimilis,
1150, 1151, 1212.
— virgatus, 1151,
1212.

virgatus, 1151.

— virgatus, 1151.
Hymenolepis, 652.
Hyperoodon

rostratus, 663.
Hypocnemis

neévioides, 1167.

— capnitis, 1168.
Hypolimnas

bolina, 702, '725, 735,

737, 738, 740.
diocippus, 727, 739.
misippus, 702, 703,

726, 727, 737, 738,

739, 740, 744.

Hypopyrrhus

pyrohypogaster, 1123,
Hypothymis

azurea, 721.
Hypsipetes

ganeesa, 717.

olivaceus, 700.
Hypuroptila

buffoni, 1185.

urochrysa, 1184.

784,

XXVill

Icterus
chrysocephalus, 860.
Jamaicai, 860.
mesomelas, 1123.

— taczanowskit,
1128.
salvinit, 1123,
vulgares, 860.
Ictidognathus, gen. nov.,
1078.
parvidens, 1078, 1081,
1082.

Ictodosaurus, 1079.
angusticeps, 1078.

Idiogenes, 652.

Tole
icterica, 717.

Irena
puella, 716.

Tthomia,
zelica, 705.

Jainides
bochus, 729.
Jonornis
martinica, 1208.
Juncella
elongata, 888.
flagellum, 838.
spiralis, 871, 889.
Junonia
lemonias, 727.
(Precis) almana, 738.

Kallima, 727.
philarchus, 728.

Kelaartia
penicillata, 717.

Lacerta
agilis, 864.
dugesit, 864.
muralis, 864.
— fjilfolensis, 864.
ocellata, 837.
viridis, 864.
vivipara, 837.
Lalage
maculosa, "Td4.
rufiventer, 699, TOL.
(Oxynotus) newton,
698, 699.
Lampides
batica, 701.
Lanius
collurio, 859.
cristatus, 719, 737,
739.

Lanius
erythronotus, 719.
lineatus, 1157.
nevius, 1158.
Laphyetes
satrapa, 1138.
Larvivora
brunnea, 716.
Lateriporus, 652.
Lathria
unirufa, 1144.
— castaneotincta, 1144
1145, 1212.
— clara, 1145, 1212.
Lechriorchis
elongatus, 681.
primus, 681.
validus, 677, 681, 682,
685, 685, 686,

?

(Renifer) — edongatus,
677.

Legatus

albicollis _albicollis,
1134.

Lemur

coronatus, 856.

Julvus albifrons, 856.

macaco, 856, 994,

1003.

mongoz, 850.

varius, 671.
Leontocebus

edipus, 856.

rosalia, 856.

rufimanus, 806.

Leptocheirus

aberrans, 561, 562,
563.

bispinosus, 562, 563,

572, 576, 585, 586,
587, 589, 594.
cornuauret, 561, 562,
563, 564, 569.
dellavallet, 561, 568,
576, 577, 582.
fasciatus, 576, 585.
guttatus, 561, 562,
563, 572, 576, 585,
587, 588, 589, 593.
hirsutimanus, 561,

563, 571, 588, 594.

massiliensis, 561,
589.
pectinatus, 561, 563,

572, 573, 574, 575,
576, 577, 582, 584,
585, 594.

ptiosus, 361, 562, 563,
564, 569, 57 0, ome
573, 576, 584, 585,
594.

INDEX OF SCIENTIFIC NAMES.

Leptocheirus
pm. ae 561, 562, Bee,
ee. 561, 562,

563, 564, 569. )
tricristatus, 561, ob?
572, 573.
Leptodira
annulata, 681.
Leptodon
cayennensis, 1205,
palliatus, 1204.
Leptopogon
poliocephalus, 1182.
superciliaris, 1132.
— poliocephalus, 1152.
— supercilaris, 1132.
— transandinus, 1182.
Lepus
jilchnert, 692.
swinhoei, 692.
— brevinasus, 692.
Lethe
daretis, 709, 710, 735,
737.
Leucolepis
pheocephalus lawrencii,
1089, 1213.
— pheocephalus, 1088,
1089, 1212.
Leucopternis
plumbea, 1204, 1210.
schistacea, 1204.
semiplumbea, 1204.
Linax, 804.
agrestis, 814.
arborium, 814.
maximus, 813.
Limenitis
sthylla, 865.
Limnoenida, 1046, 1047,
1049, 10538.
tanganice, 1056.
Linstowia, 636, 637.
Liothrix
luteus, 8d8.

Lipaugus
holerythrus, 1145,
1146.
— holerythrus, 1146.
1212.
— rosenbergi, 1145,

1146, 1212.
Lizzia, 1046.
Locusta

viridissima, 835.
Lonchodes sp., 839.

Lophiomys
ibeanus, 946, 947.
imhaust, O47,
948.

Loxia
grossa, 1121.
Loxoconcha
decipiens,
601.
impressa, 595.
obesa, 595, 597, 601.
‘subalata, 593, 598,
601
Lucanus
cervus, 839.

Lucidella, 761, 773, 776,
His, TD, TO, Wea,
729, 800.

aureola, 762, '763, 772,
780, 786, 793, 807,
808.

Lutreola

sibirica, 688.

Lycena

astrarche, 830.

icarus, 830.

595; (97,

(Zesius) chrysomellus, |

739, 740.
Lycosaurus, L078.
curvimola, 1079.
pardalis, 1079.
tigrinus, 893, 1079.
Lygosoma, 705, 706,
Lyriocephalus
seutatus, 700.
Lystrosaurus, 1080.

Mabuia, 705.
carinata, 706.
Macacus
cynomolgus, 651.
pileatus, 856.
radiatus, 650.
Macrocypris
decora, 595.
Macroglossa sp., 732.
Macropteryx
coronata, 723.
Macropus, 918, 938, 940.
bennetti, 927.
billardieri, 926.
brachyurus, 927.
eugentt, 927.
giganteus, 927, 939.
Macrorhinus
crozetensis, 671.
Madoqua
damarensis, 977.
hararensis, 979.
phillipsi, 978-984.
— gubanensis, 979,
983, 984.
— hararensis, 988,
984

INDEX OF SCIENTIFIC NAMES.

Madoqua

piacentinit, 981, 984,
swaynet, 980, 981, 982,
984.
Malacogorgia
capensis, 871, 885.
Malacoptila
mysticalis, 1197.
panamensis, 1196, 1197.

—- panamensis, 1197,
1212.

— poliopis, 1196, 1197,
IPARS

poliopis, 1196.
Malleus, 1060.
Mallotus

villosus, 666.
Mainestra

oleracea, 834.

persicarié, 832, 834,

835.
Mantis

religiosa, TOL.
Margaritifera

margaritifera, O57,

1058, 1059, 1060,
1061.
mauritit, 1052, 1060.
vulgaris, 1059, 1060,
1065.
Marmaronetta

angustirostris, 558.
Masius

chrysopterus _ bellus,

1138, 1139, 1211.

— coronulatus, 1139,

1211.

coronulatus, 1138.
Megacephalon

maleo, 757.
Megalama

virens, 862.
Megalobatrachus

maximus, 869.
Megapodius, 749, 750.

andersont, 758.

brazieri, 751.

brenchleyt, 751, 752,

cuming?, 756, 757.

duperreyt, 751, 753.

eremita, 752, Td8.

freycineti, 752, 758.

geelvinkianus, 757.

huttont, 756.

laperousit, 757.

layardi, 757.

macgillivrayt, 751.

pritchardi, "754, 755,

756, 757.

senex, 753, 757.

wallacei, 751.

XX1X

Megaptera
boops, 662.
Meiglyptes
loricatus, 1188.

- Melanargia

galathea, 816, 824, 827,
828, 865, 867.

Melanerpes
pucheramt perileucus,
1188.
— pucherani, 1188.
Melanitis

leda, 747.
tambra, TAT.
Meleagris
americana, 863.
ecellata, 863.
Melibe
jfinbriata, 1064, 1065.
Melierax
monogrammicus, 869.
Melinodon, 916, 923.
simus, 913, 923, 925.
Melita
artemis, 815, 820. 824,
825, 826, 829, 831,
866.
athalia, 866.
aurinia, 825, 866.
cinxia, 866.
Melitodes, 890.
dichotoma, 870, 877.
espert. 870, 874.
nodosa, 870, 876.
Melolontha
vulgaris, 839.
Meriones
crassus, 986.
Merops
ornatus, 869.
philippinus, 722.
swinhoei, 722, 733.

viridis, 722, %27,
732,
Merula

kinnist, 722.

tristis, LO86.
Mesopicos

eecilti, 1188.
Metroliasthes, 637.
Micrastur

guerilla, 1208.

— interstes, 1203.

— zonothorax, 1203.

interstes, 1203.

ruficollis, 1203.
Microconodon, 924.
Micromys

minutus, 690.
Micropogon

bourcierti, 1199.

XXX

Microtus

calamorum, 691.

— superus, 691.

johannes, 687, 691.

mandarinus, 691.

nux, 687.

(Caryomys)

692.

(—) nue, 691.
Milax

sowerby?, 814.
Millepora

alcicornis, 1058, 1060.
Mimus

orpheus, 858.

polyglottus, 858.

saturninus, 858.
Miniopterus, 910, 920.
Mionectes

oleagineus hederaceus,

1131.
olivaceus, 1131.
— galhinus, 1131.

eva,

— hederaceus, 1181,
1210, 1212.

— olivaceus, 1131,
1212.

— venezuelensis, 1131.
striaticollis, 1131.
— poliocephalus,
1131.
Mitrospingus
cassinit, 1116.
Mniotilta
varia, 1091.
Merrisia, 1048, 1050.

lyonst, 1045, 1046,
1047, 1049, 1050,
1058, 1054, 1055,
1056.

Molpastes

hemorrhous, 717.

Momotus

equatorialis, 1194.
— equatorialis, 1194.
— chlorolemus, 1194.
lessont, 1194.
martit, 1193.
semirufa, 1193,
subrifescens, 862.
Monasa
mystacalis, 1197.
pallescens, 1197, 1198,
1211.
Monopylidium, 652.
Mosvhops, gen. nov.,
1073

capensis, 1078, 1082.
Motacilla, 698.
estiva, 1091.

Motacilla
cayana, 1095.
noveboracensis, 1091.
ruticilla, 1098.
spiza, 1096.
varia, 1091.

Mungos
albicauda, 856.
Julvescens, 856.
galera, 857.
mungo, 856.

Murex, 1060, 1061.
ramosus, 1057, 1059.

Muriceides, 880.
Jusca, 870,

893.

Mus
decumanus, 945,
wagnert, 690.

Muscicapa
cayanensis, 1134.
Fulvicauda, 1092.
purpurata, 1148.

Muscipeta
borbonica, 701.
ralloides, 1087, 1088.

Myadestes
ralloides, 1087.

Mycalesis, 710.
ceyloniea, 728, 740,

TAL.
narcissus, 747.
Mycetes

878,

niger, 656.
Myiarchus

crinitus, 1137.

nigriceps, 1137.

tuberculifer, 1137.

tyrannulus, 1137.

Myiobius

barbatus, 1135.

— atricaudus,
1136.

— barbatus, 1136,

— mastacalis, 1135,
1136.

erythrurus, 1136.

— fulvigularis, 1136.

fulvigularis, 1136.

mastacalis, 1125.

sulphureipygius, 1135.

— aureatus, 1135,
1136, 1212.

— sulphureipygius,
1136.

1135,

— villosus, 1135, 1136,
1212.

villosus, 1185.

xvanthopygus, 1135.

— aureatus, 1135.

INDEX OF SCIENTIFIC NAMES.

Myiochanes
richardsonit
soni, 1136.
Myiodioctes
tristriatus, 1091.
Myiozetetes
eayanensis cayanensis,
1134.
— erythroptera, 1134.
rufipennis, 1134.
texensis, 1134.

richard-

| Myothera

analis, 1173.
Myotis
myosotis ancilla, 687,
688.
Myrmeciza
berlepschi, 1168.
exsul, 1169.
leucaspis, 1172.
zeledoni, 1169.

Myrmelastes
cassini, 1169, 1170.
exsul exsul, 1170,
1212.
— maculifer, 1169,

1170, 1212.
tmmaculatus, 1169.
— berlepschi, 1169.
— immaculatus, 1168.
— zeledoni, 1168.

Myrmothera
axillaris, 1162.
Myrmotherula
axillaris melena, 1162.
cineretventris, 1163.
fulviventris, 1162.

— fulviventris, 1162,
1212.

— viduata, 1162,
1212.

melena, 1162.

menetriestz, 1163.

— schisticolor, 1163.

ornata, 1162.

sancte-marte, 1163.

schisticolor, 1163.

— sancte-marte, 1163.

— schisticolor, 1163.

surinamensis, 1160,
1161.

— multostriata, 1159,

1159,

— surinamenis, 1159,
1160.
vidua, 1162.
Myzantha
garrula, 860.

INDEX OF SCIENTIFIC NAMES.

Navicella, 784.
Nematoteenia, 637.
Nemeobius
lucina, 831.
Nemorhzdus sp., 695.
Nemosia
rosenhergi,

1116,

. |
Neoniphargus, 949, 950, |
951.

Neptis
leucothoé, 735, 736.
lucilia, 865.
sapphe, 865.
varmona, 721.
Nerita, 764, 766.
Neritina, 762, 764.

Niphargus, 949, 950,
951.

Noctua, 818, 819,
835.

Nosema

apis, 625, 626.
bombyeis, 625.
Nothocercus
bonapartii, 1208.
Jrantzti, 1208, 1213.
intercedens, 1208,
1211, 1213.
Nyctipithecus
trivirgatus, 856.
Nythosaurus, 900, 901,
902, 905, 907, 908,
915, 919, 928.
larvatus, 894, 898,
899.

Obelia,
1355.
Oceania, 1049.
Ochetosoma
Jormosum,
686.
menstruosum, 681.
Ochotona
cansa, 692, 6935.
sorella, 692.
syrinx, 692.
Ocypode, 1066.
Ocypus
olens, 812, 816, 8836,
837, 838, 846, 855.
Odontophorus
cinctus, 1206, 1207.
parambe baliolus,
1211.
— parambe, 1211,
spodiostethus, 1206,
1207.

1046, 1048,

681, 683,

Ckdicnemus
bistriatus, 864.
Oligorchis, 652.
Oochoristica, 627, 630,
631, 632, 635, 6387,
654, 655, 658, 1000,
1005, 1017, 1018.
sp., 627, 628, 633.
rostellata, 1018.
tetragonocephala, 629,
638.
wageneri, 638.
Ophion
luteus, 848.
Ophisaurus
apus, 864.
Opisthoctenodon, 1075.
Orchistoma
pileus, 1051.
Oreocincla
imbricata, 722.
spiloptiera, 722.

| Oreotragus

saltator, 269.
— porteust, 960.
Oriolus
maculatus, 859.
oryzivorus, 1122.
Ornismya
kingtt, 1187.
Ornithoptera
darsius. 712.
Ornithorhynchus, 908,
910, 918, 919, 920,
921, 922, 942.
Orobophana, 761, 762,
773, 788, 726, 797,
798, 799, 800.
ponsonbyt, 772, 778,
780, 782, 783, 793,
807, 808, 809.
Ortalida
goudotii, 1207.
Ortalis
ruficauda, 863.
Orthogonys
olivuceus, 1114.
Orycteropus, 921.
Oryzoborus
ethiops, 1098.
funereus, 1098.
— ethiops, 1098.
— funereus, 1098.
Ostinops
_ atrocastaneus, 1211.
salmoni, 1211.
viridis, 860.

Ostracolethe, 804.
Ostrea sp., 1059,
1060.

XXX1

Otaria

californiana, 985.
Otis

ludwigi, 863.

vigorsti, 863.
Otocompsa

flaviventris, 858.
Otocryptis

bivitiata, 706.
Oudenodon, 1080.

bolorhinus, 1076, 1081,

1082.

strigiceps, 1076.
Ourebia

nuigricaudata, 869.
Oxynotus

JSerrugineus, 701.

Pachyrhamphus
atricapillus, 1144.
castaneus, 1144.
cinereiventris, 1143,
cinnamomenus, 1144.
dorsalis, 1143, 114,

1211.
marginatus, 1144.
niger cinereiventris,
1144.
ru,fescens, 1144.
rufus, 1144.

Paguma
larvata, 688.

Paleohelicina, 761, 762,

772, 113, 782, 787,
796, 797.

ide, 763, 770, 778,
780, 793, 807, 808,
809.

Panoplites
flavescens, 1185.

Pantodon
buchholzi, 985.

Papilio
agamemnon, 730.
aristolochie, 702, 709,

741, 748, 744.
clytia dissimilis, 702,
703, 705.

— lankeswara, 702,
704.

— panope, 705.

demodocus, 748.

demoleus, 08, 27,
748.

epiphorbas, 702.

hector, 702, 713, 714,
72%, Tdi, 40, 741,
744.

jason, 729.

exon
Papilio
lankeswara, 702, 704,
7A0.
limniace, 70).
machaon, 730.
manlius, 702.
phorbanta, 698, 700.
polytes, 702, 707, 708,
713, 714, 719, 729,
732, 743.
— romulus, 702.
surpedon, 727, 734.
(Menelaides) hector,
708, 709, 740, 743.
Papio
porcarius, 558, 559.
— griseipes, &58,
559.
sphinx, 842, 856.
Paracrangonyx, 949,
950.
compactus, 952.
Paracyamus
boopis, 668.
Paracyathus
cavatus, 1019, 1027,
1028, 1039, 1044.
erassus, 1039.
Paradoxostoma
arcuatum, 595.
cylindricum, 595.
flexuosum, 595.
_ gracile, 595, 600, 601.
- hibernicum, 595.
Paradoxurus
larvatus, 621.
Paranerita, 766,
779.
gagates, 764, 765.
Pareiasaurus, 919, 1079,
1080, 1081.
Pareronia
ceylonica, 702.
Parlotichus, 919, 920.
Parus
major, 858.
Paruterina, 652.
Passer
domesticus, 698.
Pellorneum
Suscicapillum, 715.
Pennella, 667, 668.
Penthoceryx
sonneratis, 724.
Perameles, 939, 942,
Perarge
megera, 815, 826,
Pericrocotus
flammeus, 719.
peregrinus, 719.

178,

' Phalanger, 1017.

Perisoreus
canadensis, 861,

Petanrista
alborufius, 689.

Petrogale, 940.

penicillata, 939.
Phethornis

berlepschi, 1179, ©

1180.

columbianus, 1179. °

syrmatophorus, 1179,
1180. i

— columbianus, 1180.

— syrmatophorus, °
1179, 1180,

yarugui, 1178.

— sancti-johanms,
1178, 1179, 1211.

— yaruqui, 1179,
PAD

Phalangista, 1017.
Phalangium sp., 814.
Phalera

bucephala, 832.
Phascolarctos, 1017.
Phasianus

principalis, 862.

reevesit, 862.
Pheasia

dicted, 833, 835.

tremule, 833, 834.
Phedina

borhonica, 698, 699.
Philydor

philydor

1151.

virgatus, 1151.
Pheenicothraupis

cristata, 1118, 1114.
Phrygilus

aldunatit, 558.
Phyllades

consolisma, 730.
Physeter

macrocephalus, 663.
Picumnus

canus, 1189, 1190.

dimotus, 1192.

granadensis,

1190, 1191.
olivaceus, 1189, 1190,

virgatus,

1189,

1192.

— dimotus, 1192.

— flavotinctus, 1190,
1191.

— granadensis, 1189,
1190, 1191, 1211,
1212.

— harterti, 1190, 1191,

211.

INDEX OF SCIENTIFIC NAMES.

Picumnus
' olivaceus _ olivaceus,
1190, 1191, 1192.
— panamensis, 1190,
Zale
Picus

lineatus, 1189.
rubiginosus, 1187.
(Chloropicus) iri,
1188.
_ Pieris
brassice, 816, 818, 823,
829, 834, 835, 846,
864.
napi, 815, 826, 827,
828, 829, 846, 865.
rapé, 815, 820, 826,
835, 865.
_ Pimelepturus
cinerescens, 1065.
Pionopsitta
hematotis, 1202, 1213.
pulchra, 1202, 1210,
1213.
Pionus
menstrius, 1102.
Pipra
chrysoptera, 1138:
coronata, 1140.
— velutina, 1140.
cyaneocapilla, 1140.
mentalis, 1139.
— ignifera, 1139, 1212.
— nmunor, 1139, 1140,
1212.
velutina, 1140.
vitellina, 1141.
Piranga
rubra rubra, 1113.
Pitangus
Ubovittatus, 1134,
parvus, 1135.
. sulphuratus, 861.
Pithys
bicolor, 1170, 1173.
— equatorialis, 1172.
— olivascens, 1173.
leucaspis, 1172, 1173.
Pittasoma
michleri, 1212.
rosenbergt, 1175, 1176,
1211, 1212.
rufopileatum, 1176,
1211.
Pitylus
grossus, 1121.
Planorbis, 762.
Platypsaris
homochrous, 11438.
Platyrhynchos
olivaceus, 1127.

INDEX OF SCIENTIFIC NAMES,

Platyrhynchos
virescens, 1136.
Platytriccus
albogularis, 1125,
1126.
cancroma, 1125.
insularts, 1126.
mystaceus, 1125.
— albogularis, 1125,
1126.
— insularis, 1126
— mystaceus, 1126.
Pleurotomaria, 803, 804,
805.
Plocamopherus
ocellatus, 1064.
Pneumatophilus, 683.
Podocoryne, 1046, 1047,
1049.
carnea, 1058.
Poéphagus
grunniens, 869.
Polydelphis, 620.
Polyerata
reint, 1181, 1182.
vosenbergi, 1181, 1182,
1210.
— reini, 1181.
Polyommatus
betica. 709, 740.
Polytes, 744.
Pomatorhinus
melanurus, 715.
Pontocypris
dispar, 595.
succinea, 595.
Porphyrio
melanonotus, 980,
martinicus, 1208.
Porzana
albigularis, 1208.
Pratincola
atrata, 721.
(Motacilla)
698.
Precis
almana, 740.
lemonias, 737.
rhadama, 747.
Prinia
Jerdoni, 719.
socialis, 718.
sylvatica, 718.
Prioneris
sita, 702,
704.
Prionites
martit, 1198.
Priotrochatella, 761.
risterodon, 1075.

sybilla,

703,

Procavia
dorsalis, 673.
emini, 673.

Procnias
cerulea

1099.

occidentalis, 1099.

tersa, 1099.

Eres opnen: 919, 920,

922, 1080.

Prodicynodon, 1076.

Progne
chalybea, 1093.

— chalybea, 1093.
Progynia, 652.
Proorchida, 652.
Propappus, 1080.

occidentalis,

Proserpina, 760, 761,
762, 799.
Protomedeia
fasciata, 568, 576,
585.
guttata, 561, 572, 573,
576.

hirsutimana, 561, 56
568, 571, 576, 58
589.

— massiliensis, 562.

pectinata, 576, 584.

pilosa, 563, 576, 577
585.

Psammogorgia, 883.

pulchra, 871, 881, 892,
893.

Psarocolius

mesomelas, 1122.

Pseudoclytia
pentata, 1048.
Pseudotrochatella
undulata, 761.
Psittacus
menstruus, 1202.
Psittospiza

riefferii, 1120.

—— elegans, 1120.

— riefferii, 1120.

Psophia

crepitans, 864.

Pteroglossus
erythropygius,
1202

ato

12018

— erythropygius, 1201,
1202, 1211.

— sanguineus, 1201,
1202, 1211.

sanguineus, 1201.

Pterostichus, 836,

855.

niger, 816, 837, 842,

846.

(Abax) striola, 837.

Proc, Zoou, Sov.—191L, No, LXX XIII,

XXX111

Pterostichus
(Steropus) madidus,
816, 838, 842.
Ptilocheirus
pectinatus, 576, 585.
tricristatus, 563,
576.
Ptilotis
carunculata, 754.
Putorius
nigripes, 097, 559.
Pyenonotus
hemorrhous, 858.
Jocosus, 701.
leucogenys, 808.
leuconotus, 858.
luteolus, (17.
melanicterus, 717.
rzanthopygus, 858.
Pyctorhis
nasalis, 715.
Pyrameis
atlanta, 823.
cardut, 709, 710,
738, 823.
Pyramidula, 803.
Pyranga
estiva, 1115.
Pyrgoma
stokesii, 1029.
Pyriglena
berlepschi, 1167.
maculicaudis, 1166.
tyrannina, 1165.
Pyrophyllia, 1088, 1041.
inflata, 1018, 1019,
1020, 1037, 1039,
1040, 1044.
Pyxis
arachnoides, 624.

O72,

736,

Querquedula
cyanoptera, 1209.
Querula
eruenta, 1148.
purpurata, 1148.
Quiscalus

gundlachi, 671.

Rallus
abbotti, 864.
Ramphocelus
icteronotus,
1113.
Rathkea, 1046, 1047.
Renifer, 677, 685.
ellipticus, 683.
sauronates, 681, 685.
Retorquata, 761.
83

1112,

XXXIV

Rhabdias
bufonis, 620.
Rhabdometra, 637.
RKhagonyche, 828.
Julva, 812, 840, 842,
850.
Rhamphastos
ambiguus, 1201.
swainsonit, 1200, 1201.
tocard, 1200.
tocardus, 1201.
Rhamphoeenus
cinereiventris, 1164.
— cinereiventris, 1164,
1212.
— semitorquatus, 1164.
semitorquatus, 1164,
1212.
Rhamphocelus
brasilius, 860.
icteronotus, 1112.
Rhinoceros, 945.
bicornis, 959, 985:
somaliensis, 9059,
960.
Rhinochetus
Jubatus, 864.
Rhbinolophus
Jerrum-equinum, 687.
Rbipidura
albifrontata, 721.
tricolor, 859.
Rhopocichla
nigrifrons, 715.
Rhopoctites
alogus, 1149.
Rhynchoeyclus
equinoctialis, 1 127,
cinereiceps, 1126.
— cinereiceps, 1212.
— flavotectus, 1126,
1127, 1210, 1212.
meg actphala, 1126.
— flavotectus, 1126.
sulphurescens, 1127.
— asemus, 1127.

Rhynchortyx
cinctus, 1206, 1207.
Rhynchotragus, 977,
978.

cordeauat, 983.

erlangeri, 983, 984.

guentheri, 980-984.

— wroughtont, 983,

984.

kirkii, 978, 982.
Rictularia

plagiostoma, 620.
Rivulus

flahellicauda, 985.

ocellatus, 985.

Rivulus
poeyi, 98d.

Salmo

salar, 671.
Saltator

atripennis, 1120.

elegans, 1120.

magnus, 1121.

maximus, 1121.
Sapayoa

enigma, 1141,

1213.

Saysia, 1046, 1056.

prolifera, 1066.
Sarsiella

capsula, 595.
Satyrus

semele, 830.
Saurosternon, 1080.
Sayornis

ardesiacus, 1125.

cineracea, 1125.

— cineracea, 1125.
Schcenicola

platyura, 718.
Scirpearella sp., 888.

aurantiaca, 888.

indica, 888.
Scirpearia sp., 888.

flagellum, 871, 888,

3g.

furcata, 871, 888.

ochracea, 839.
Sciurotamias

davidianus, 689.
Sclerochilus

contortus, 595.

levis, 695, 601.
Scops

leucotis, 862.
Scotothorus, 1141.
Scylacosaurus, 919.

constrictus, 1078.
Scymnosaurus sp., 1081.
Seiurus

noveboracensis, 1091.

— noveboracensis, 1091.
Semnornis

ramphastinus, 1200.
Septaria, 764, 760, 784.
Serpophaga

albagrisea, 1183.

cinerea, 1130.

— cana, 11380.

— grisea, 1130.

grisea, 1130.

parambe, 1138.
Sesamodon, 909,

923.

1210,

916,

INDEX OF SCIENTIFIC NAMES.

|
}

Sesamodon
browni, 913, 914, 915,
923, 925.
Setophaga
ruticilla, 1093.
Sialia
sialis, 857.
Sibia
capistrata, 859.
Siderastreea
radians, 1084.
Siphia
hyperythra, 720, 730.
Siptornis
erythrops erythrops,
1149, 1211.
— griseigularis, 1149,
1211, 1212:
— rufigenis, 1149, 1212.
Sitta
surinamensis, 1160.
Siurus
noveboracensis, 1091.
Spermophila
ophihalmica, 1088.
Sphecodes, 867.
Spheniscus
demersus, 571.
Sphenodon, 922.
Sphenuta
subulata, 1150.
Spilesoma
melanopsis, 709,
Spondylus, 1059.
sp., LOL.
Spongioderina, 872.
verrucosum, 870, 87-4.
Sporophila
aurita, 1099, 1212.
hicksii, 1099.
ophthalmica,
1099, 1212.
Stachvodes, 891.
gichristi, 871, 888,
893.
trilepis, 886.
Stelgidopteryx
ruficollis, 1093.
— uropygialis, 1098.
wropygialis, 1093.
Stenobothrus sp.,
846.
Stephanoseris
rousseaut. 1022, 1026,
Stilesia, 1002.
Stomotoca
dinema, 1051.
Stoparola
sordida, 720:
Strangalia
armatt, 839.

1098,

835,

muelanocephala, C39, |

650. {

Tetragonops
ramphastinus, 1200.

INDEX OF SCIENTIFIC NAMES. XXXV
Struthidea | Tenia | Tetrao
cinerea, 671, 859. mucronata, 649, australis, TAS.
Struthio tetragonocephala, 637. Thalassochelys
australis, 987. tuberculata, 635. | caretta, 624.
massaicus, 987. (Leniarhynehus) sagi- | Thalurania
molybdophanes, 987. | nata, 1018. funnyt, 1183, 1184,
Sturanya, 761. | Tamandua | 1,
Sturmopastor | tetradactyla, 636, 637, —_verticeps, 1184, 1211.
contra, 860. 638, 1018. Thamnobia
julle, 860. | Tanagara fulicata, 721.
Sturnornis | notahitis, 1111, Thamnophilus
senex, 720. | Tanagra atrinucha, L158.
Sturnus | maxima, 112). cachabiensis, 1167.
holosericeus, 1122, | melanoptera, 1112. immaculatus, 1163.
Stylocyathus, 1050. | palmarum., 1112. major, 1157.
Suastus — melanoptera, 1112. — granadensis, 1158,
gremius, 732, — vjolilavata, 1112. — melanurus, 115%.
Suberia, 872. peruviana, 1104, — transandeanus,
capensis, &70, 871, 873, | riefferti, 1120. 1157, 1158.
fohS)ZF, teh )5}, ruficer via, 1108. nevis, 1158.
clavaria, 873. (Aglaia) aurulenta, — atrinucha, 1158.
gentht, 873. 1102: | — nevius, 1159.
hellikeri, 873. (—) labradorides, 1109. | qurwensis, 1168.
Sulphurina, 761. (Calliste) ryfiverter, | fransandeanus, 1157,
Suricata 1108. | 1158.
suricatta, 856. | Tanagrella | Thanaos
~ Surniculus rufigula, 1102. tuges, 825, 831, 848.
lugubris, 724. | Taognathus, gen. nov., Therapha
Sus sp., 693. 1076. | — hyocyami, 846, 847.
Sylvia megalodon, 1076, 1081, | Vherjodontia, 894,
castanea, 1091. 1082. | Lhouarella
Symphalangus | Tapinocephalus, 919, | hicksoni, 886, 893,
syndactylus, 671, | 1074. striata, 871,
Synallaxis | Telehinia | Threnetes
castanea, 1148. viole, 704, 708, 709, | fraseri, 1177.
erythrops, 1149, G4, 726, 730; 740) | puckeri, 1177.
pudica, 1148. 7 ae 741. — fraser’, V77, W212)
rugigents, 1149. | Temenuchus — Tinea, — MAl7T
uniriufa, 1148. | pagodarumn, 720. 1212.
Synictis Vephrodornis Thrinaxodon, 899,
penicillata, 856, pondicerianus, 719. Thripadectes
Syrnium Teracolus flammulatus, 1149,
nuchale, 869. danae, 713. selateri, 1149, 1211.
Syrphus, 850. Jausta, 713, 727. Thryophilus
Terias leucapogon, 1089, 1090
Tachornis jfloricola, TAT. DUD
batassiensis, 723. hecabe, 708, 709, 710, nigricupillus, 1089.
Tachyphonus 727, 730, 735=747. — nigricapillus, 1089,
cassinit, 1116. libythea, 709, 735, ae
chrysomelas, 1117, Terpsiphone | — schattii, 1089, 1211,
delutrii, 1115, 1116. paradisi, 721. PA
melaleucus, 860. Tersina schotizi, 1089.
propingwus, 1115. viridis oceidcntalis, semibadius, 1089,
rubrifrons, 1115. 1099, NB,
xanthopygius, 1114, Testudo, 945. thoracicus, 1090, 1212,
1115. daudinii, 623. Thryothorus
Tenia, 994, 1003, 1017, elephantina, 628, | nigricapillus, 1089,
1018. greca, 624. | Thysanocephaluu
conferta, 6A9. nigra, 624, | crispum, 629.

Thysanosoma, 994, 996,
1000,

XXXV1

Thysanosoma
gambianum, 651, 652,
653, 658, 659, 995,
1002, 1008.
Thysanotenia, gen. nov.,
994, 1002.
gambiana, 997-1000,
1002, 1003.
lemuris, 994-1008,
1008.
Timarcha, 842.
tenebricosa, 819, 837,
841, 843, 845.
Tinamus
bonapartei, 1208.
Jrantzti, 1208.
Tinnunculus
alaudarius, 724.
punctatus, 700.
Tipula
oleracea, 848.
Tityra
albitorques, 1142.
— albitorques, 1142.
buckleyi, 1148.
personata, 1142.
semifasciata, 1142.
— columbiana, 1142,
1212.

— costaricensis, 1142, |

1212.
— semifasciata, 1142.
Todirostrum :

cinereum, 1128.

— cinerewm, 1128.

— sclateri, 1128.
Todus

cinereus, 1128.
Trachyphonus

caffer, 862.
Trematotrochus, 1038.

Jenestratus, 1030.

verconis, 1080, 1039.

zelandie, 1019, 1620,
1029, 1081-1084.
Treveleyana
erocea, 1064, 1068.
Tribonyx
ventralis, 864.
Triccus

sclateri, 1128.
Trichocephalus, 674.
Trichogaster

lahius, 985.
Trichoglossus

nove-hollandie, 671.
Trichosoma, 674, 676,
Trichostrongylus

retorteformis. 676.
Trichosurus, 918.
Trichuris, 674.

INDEX OF SCIENTIFIC NAMES.

Tridaena, 1059. |
gigas, 1058, |
Trionyx, 945, |
Trirachodon, 894, 905, |
908, 909, 913, 923. |
hannemeyeri, 906,
907.
Tritylodon, 898, 918,
919.

longevus, 925.
Trochalopteron
canorum, 859.
erythrocephalum, 859.
nigrimentum, 859.
Trochatella
chrysochasma, 798. |
Trochilus
aquila, 1180. |
aurelie, 1185. |
barroti, 1186. |
benjaming, 1186.
exortis, 1186.
Fannyi, 11838.
Navescens, 1185.
Jrancie, 1182.
hirsutus, 1178.
mellivorus. 1181. |
vuckeri, 1177.
feacatl, 1182.
yaruqui, 1178.

Trochocerecus
borbonicus, 698, 699,
7Ol.

Trochosmilia, 1040.
Trochus, 794.
Trogon
macrurus, 1193,
massena, 1193.
melanurus macrourus,
1195.
Trogopterus
xzanthipes, 689.
Tropicoris
rufipes, 840, 847.
Tropidonotus
rhombifer, 688.
Trypanosoma
lewis, 945.
Turbinolia, 1020, 1030.
Turbo, 794.
Turdus
ertnitus, 1137.
dague, 1086.
musicus, 857.
swatnsenii, 1087.
tristis cnephosa, 1087,
1212.
— dague, 1086, 1087,
1212:
ustulatus, 1087.
Turritopsis, 1049,

Tyrannula
ardosiaca, 1125.
cineracea, 1125,
erythroptera, 1134.
richardsonti, 1186.
sulphureipygius, 1135.

| Tyrannus

albicollis, 1134.
melancholicus, 1138.
— satrapa, 11388.

Udenodon, 923.
Upupa
epops, 862,
Uranomitra
Francie, 1182.
Urospatha
martit, 1198.
—— martii, 1193.
— semirufa, 1198.
Urosticte
benjamini, 1186.

| Valvata, 762.

Vanessa
haronica, 735.
Zo, 812, 822, 865.
urtice, 820,
865.
Veniliornis
kirkii cecilii, 1188.
— dariensis, 1188.
Viminella
Hagellum, 888.
Vincta sp., 748.
Viverra
jilchneri, 688.
etbetha, 688.
Volucella
bombylans, 812, 847,
853. 854, 855, 867.
Vulpes sp., 688.

822,

Waldemaria
Japonica, 784.

Xanthornus
mesomelas salvinii,
1122, 1128.
— taczanowshii, 1128.
Xenicopsis
mentalis, 1151, 1152.

subalaris lineatus,
1151.
— subalaris, 1151.
Xenops

genibarbis, 1152.

INDEX OF SCIENTIC NAMES. XXXVI1

Xenops Xestoleberis ; Zesius
genibarbis littovalis, nigromaculata, 595, | chrysomallus, 727.
1152; W212. 598, 6OL. Zizera, 722.
mexicanus, 1152, | Xiphophorus Zosterops
1212. helleri, 985. ceylouensis, 716.
littoralis, 1152. Xiphorhynchus chloronota, 701.
mecvicanus, 1152 lacrymosus rostratus, hesitata, 698.
ruficauda, 1152. 1153. mauritiana, TOL.
Xenopus, 919. palpebrosa, 716.
Xestoleberis Yyhthima, 710. (Malacirops) borbonica,
depressa, 595. 698.
latissima, 595, 598, GOL. | Zebrapicus Zschokkeella, 651, 660.

maryar ited, I95. | pucherant, 1188. Zygeena, 867.

INDEX

OF

ILLUSTRATIONS.

Aiurosaurus tenuirosiris, Pl, UXII1.
p. 1073.
whattsi, P\. XIII. p. 1075.

Agelecyathus persicus, Fig. 220. y. 1035.

-Alcadia hollandt, Pils} XXX =—XiIL.,
p- 799.
palliata, Pls. XXX —-XXXII.,
p. 759.

Anopiotenia dasyuri, Figs. 208 215,
pp. 1004, 1006, 1007, 100%,
1010, 1012, 1018, 1015.

Aphanoconia andananica, Pl. XUL.
p. 759.
gouldiana, Pls. XXXVII., XLI.,
p. 790.

p- 799.
—— rogersti, P\. XLII. p. 759.
Argillecia affinis, Pl. XX. p. 595.
Avicula zebra, Fig. 280, p. 1058.

Bairdia dubia, Pl. XX. p. 595.
Bauria cynops, Pl. XLVI. p.
Figs. 168, 169, pp. 896, 897.
Bertiella cereopitheci, Figs.
pp. 640, 641, 643, 644, 647.
Budorcas bedfordi, P\. XX1X. p. 687.

893 ;

151-155,

Capra pyrendica hispanica, Pl. LIL.
p. 963 ; Figs. 195, 199, pp. 968,
973.

— —— pyrenaica, Bigs. 195, 198,
pp. 968, 972.

victorie, Pls. Lill, LIV.,
p- 963; Figs. 195-197, pp. 968,
970, 971.

Ceratoisis ramosa, Pl, XLIII. p. 870.

merguiensis, Pls. XXXV., XLIL, |

| Chama foliata, Pl. LX. p. 1057 ;
Fig, 229, p. 1057.
| Chromodoris inopinata, Pl. LXL
p. LOGS.

ene reticulata, P\. UXT. p. 1068.
| — ftinctoria, Pl. LXI. p. 1068.

| Columba

| Dasymetra

sp, Pls. XXIIL-XXVL.,
p- 601.

Conchoderma auritum, Big. 163, p 667.

Coronula diadema, Fig. 163, p. 667,

Cynognathus berryi, Pl, XLV E. p. 893.

-— platyceps, Figs. 171, 172, pp. 901,

903.

Cythere cingulata, Pl. XX. p. 599.

-—— crispata, Pl. XX. np. 499.

Cythereis deformis, Pl. XX. p. 595.

Cytherella ovalis, Pl. XXII. p. 595.

Cytheridets subulata, var. crenulata,
Pl XOX. p. 595:

Cytherura cribrosa, Pl, XXII. p. 598.

fossulata, Pl. XXII. p. 595.

—— maculosa, Pl. XXII. p. 595.

confenta, Pl. XOX VnLT.
p. 677.

Dendrophyllia sp., Pl. LVIT. p. 1018.

Diademodon, Pl. XLVI. p. 893.

Dielurodon whaitsi, Pl. UXIIL. p. 1073.

Diagram showing relative positions of
tooth-germs in Macropus hillardieri.

Figs. 187, 189, pp. 933, 936.

| Diagrammatic representation of two

species of corals which vary about
two distinct growth-modes, the
extremes of which converge towards
one another. Fig. 216 4, p. 1024.

representation of a single species
of coral varying about an imaginary
growth-mode. Fig. 216 2, p, L024.

INDEX OF ILLUSTRATIONS.

Dinotheriwn 1 OWI

p. 948.

hobleyi,

Equus asinus somaliensis, Fig. 202,
p. 989.

<x Hquus quagqga

chapmanni, Fig. 203, p. 992.

x Hguus zebra, Fig.

201, p. 989.

—— quagga chapmanni, Fig. 203,
p. 992.

zebra, Fig. 201, p. 989.

Eriphostoma microdon, Pl.

| p. 1073.

Eucrangonyx robertsi, Pls. XLIX.-LI1.,
p: 984.

Hunicella papillosa, Pl. XLII. p. 870.

Huplecaura media, Pl. XLV. p. 870.

Hutrochatella pulchella, Pls. XXXI.,
2LOOCUE, ZOO, XO-OOVINL, 20,
p. 799.

LXIIl.

Flabellun
p. 1018.

Play Vie

magnificum,

Gazella dorcas, Fig. 198, p. 962.

hayi, Fig. 193, p. 962.

Gomphcgnathus minor, Kigs. 175-178,
pp. 909-912.

Gorgonocephalus, Fig. 167, p. 874.

Heterocyathus equicostatus, Pl. LVIII.
p. 1018; Fig. 218, p. 1032.

alternatus, Pl. LVILTI. p. 1018.

heterocostatus, Pls. LVII., LVILII.,
p. 1018.

michelini, Fig. 218, p. 10382.

Hyla goughi, Pl. LXIV. p. 1082.

Ictidognathus parvidens, Pl. LXIII.
p. L073.

Lechriorchis validus, Pl. XXVII.
p- 677.
Leptocheirus bispinosus, Pl. XVIII.

p. 561; Fig. 146, p. 586.

guttatus, Pl. XVILI. p. 561.
—— hirsutimanus, Pl. XVIII. p. 561.
pectinatus, Pl, XIX. p. 561.
pilosus, Pl. XVII. p. 561,
pinguis, P\. XVIII. p. 561.
Lophiomys ibeanus, Fig. 190, p. 947.

XXX1X

Loxoconcha decipiens, Pl. XXT. p. 595.

obesa, Pl. XXI. p. 595.

subalata, Pl. XXI. p. 595.

LIucidella aureola, Pls. XXXII.,
XXXIV., XL, p. 759.

Macropus billardiert, Pl. XLVII. p. 926 ;
Figs. 181-189, pp. 927-929, 931-933,
9°75, 936.

Madoqua phillipsi, P\. LY. p. 977.

gubanensis, Pl, LY. p. 977.

—— —— fararensis: Pl, LY. p. 977.

—— piacentinii, Pl] LVI. p. 977.

-——— swaynei, Pl. LVI. p. 977.

Map illustrating the Distribution of
the Genus Megapodius. Fig, 166,
p- 750.

showing probable former and
present distribution of the Spanish
Ibex. Fig. 194, p. 965.

Margaritifera margaritifera, Pl. LX.
p- 1057.

Megaptera boops, Figs. 160-162, pp. 662,
664, 666.

Megapodius, Fig. 166, p. 750.

Melinodon simus, Pl. XLVI. p. 893.

Melitodes esperi, Fig. 167, p. 874.

Millepora alcicornis, Fig. 230, p 1088.

Merisia tyonsi, Pl. LIX. p. 1045; Figs.
222-228, pp. 1047, 1050, 1052, 1054,
1055.

Moschops capensis, P\. LXV. p. 1073.

Murex ramosus, Fig. 229, p. 1057.

Muriceides fusca, Pl. XLIV. p. 870.

Nythosaurus larvatus, Fig. 170, p. 899.

Ochetosoma formosum, Pl, XXVIII.
p. 677.

Oochoristica sp., Figs. 148-150, pp. 627,
628, 653.

Oreotragus saltator porteusi, Fig. 192,
p. 959.

Orohophuna pachystoma
PLL p: 759!

ponsonbyi, Pls. XXXII., XXXY.,

ponsonbyi,

XXXVIL., p. 759.

Oudenodon bolorhinus, Pl. LXIII.
p. 1078

Paleohelicina ide, Pls. XXXII.,

XXXV., XXXIX., XLI., p. 759.
Paracyathus cavatus, Pl. LVII.
p. 1018; Fig. 217, p. 1028.
| Paradoxostoma gracile, Pl, XX1.p. 590.

xl INDEX OF ILLUSTRATIONS.

Paradoxurus
p- 621.
Pigeons, Colour and Colour-pattern

Inheritaucein. Pls. XXITI.—-XXV1,
p. 601.
Porphyrio
p. 986.
Psammogorgia pulchra, Pls. XUIIL.,
XLYV., p. 870.
Putorius nigripes, Fig. 145, p. 559.
Pyrophyllia inflata, Pls. LVIL., LVIIL,,
p- 1018; Fig. 221, p. 1040.

larvatus, Fig. 147,

melanonotus, Fie. 200,

Rhinoceros bicornis, Fig. 191, p. 958.
somatiensis, Fig. 191,

p. 958.

Sclerochilus levis ?, Pl. XXII. p. 595.

Sesamodon browni, Pl. XLVI. p. 893;
Figs. 179, 180, pp. 914, 915.

Spondylus sp., Pl. LX. p. 1057.

Stachyodes gilchrist?, Pls. XLIV., XLYV.,
p. 870.

Struthidea cinerea, Fig. 164, p. 672.

= —_——_—~ —

Suberta capensis, Pls, XLIII., XLY.,
p- 870.

Taognathus LXII.
p. 1073.

Thouarella hicksoni, Pls, XLIV., XLV.,
p: 870.

Thysanosoma gambianum, Figs. 156-
159, pp. 652, 658, 658, 659.

Thysanotenia lemuris, bigs. 204-207,
pp. 995, 996, 998, 1001.

Tooth-germs in MMacropus billardieri.
Figs. 181-189, pp. 927-929, 931-
933, 935, 936.

Trematotrochus zelandie, Pls. LVII.,
LVIII., p. 1018; Figs. 218, 219,
pp. 1032, 1033.

Trichosoma sp, Fig. 165, p. 675.

Lrirachodon kannemeyeri, Figs. 173,
174, pp. 906, 907.

megalodon, Pl.

Aestoleberis latissima, Pl. XXI. p. 595.
-— nigromaculata, Pl. XXII. p, 595.

PRINTED BY TAYLOR AND FRANCIS, RED LION COURT, FLELT STREET,

PROCEEDINGS

OF THE

GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

OF THE

ZOOLOGICAL SOCIETY
OF LONDON.

1911.

PART, II.

CONTAINING Paces 557 to 868, witH 26 PuaTEs
AND 22 TEx?-FIGURES.
rs : .
J -xesonian tisty,
1" iy
7

VUI SO A9t 4

A
SEPTEMBER 1911. W/o)

PRINTED FOR THE SOCIETY,
SOLD AT THEIR HOUSE IN REGENT’S PARK.
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a [Price Twelve Shillings.

ES Ore CONT KIN es:

1911, Parr IID. (pp. 557-868).

EXHIBITIONS AND NOTICES.

ue Secretary. Report on the Additions to the Society’s Menagerie during the month of

February 1911

COE OO NON 8 Ce OO 0° OOOO DEO OOn Oh OO CHO dO Gatco noo ona oO mod OO COA mo ooUt

Mr. D. Szru-Suire, F.Z.S. Exhibition of a living Hybrid between the White-eyed
Pochard (Aythya nyroca) and the Marbled Duck (Marmaronetia angustirostris)

Mr. R. I. Pococs, F.R.S., F.L.S., F.Z.S. Exhibition of the skin of a new Chacma Baboon

(Papio porcarius, subsp. griseipes), a specimen of the North American Black-footed
Polecat (Putorius nigripes), ete. (Text-fig. 145.)

sete ee ee ee ee Oo ee ee eee eee ee te oe ee

Sir BE. Ray Layxesrer, K.C.B., F.R.S., F.Z.8. Exhibition of a special supplement to the
‘Field’ newspaper

Oe eee ee Ct Cc er we er err

Dr. R, T. Lurpzr, F.Z.8. A demonstration of Nematode parasites obtained from animals
in the Zoological Gardens

eC ee re i. rrr i, Prd

Mr. R. I. Pococs, F.R.S., F.L.S., F.Z.S8. Exhibition of a newly born Masked Palm Civet
(Paradoxurus larvatus)

eC Ce Cn CO ee i i i recy

ee ee eee eee OP ecw ee ee we we ow

Tux Secrerary. Letter on Land Tortoises in the Seychelles

Dr. H. B. Fayruan, F.Z.8., and Miss Annie Porrmr, D.Sc. Ona Bee-disease due to a
Protozoal Parasite (Nosema apis)

ee eee eee eee er PO Co eee re se eeeeee et ereesese ee thew

Tue Secrerary. Report on the Additions to the Society’s Menagerie during the month
of March 1911

ee ee ee eeco se Pe eee ese sees ee OH ee ee Cee FHP BOE ee oe PF eeseete es CO oe

Mr. C. Tare Ruean, M.A., F.Z.S. Exhibition of a series of lantern-slides of scales of the
Mallon (Sad720 Salary: ior ia oie as cale tele a tolere a wie teiotealas Cheb ce, wich chalet oleic cial aie eee

My. D. Sern-Ssirn, F.Z.S8. Exhibition of a nest of the Grey Struthidea or Apostle Bird
(Struthides cinerea), and lantern-slides of Penguins in moult and of wild Swainson’s
Lorikeets (Trichoglossus nove-hollandie). (Text-fig. 164.) ..........eseeececeeee

Dr. C. Curisty, F.Z.8. Exhibition of a collection of skins from Uganda and of a loin-cloth
taken from a native in Northern Nigeria .......... 0... 0ceeeceeeee f eievelepidereioes

Dr. Witutam Nicotn, M.A., F.Z.S. On a unique Pathological Condition in a Hare.
CMG KE tree GOS)! A26 Ne esters sate) slensieinialstauale Wiatieea fetetal choraso)o%= sic) wiehels ise sibys oh tenet ee en ee

Page

558
620
620

621
622

625
671

671

671
672

674

Contents continued on page 3 of Wrapper.

PROCEEDINGS

OF THE

GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.

EXHIBITIONS AND NOTICES.
March 21, 1911.
Dr. 8. F. Harmer, M.A., F.R.S., Vice-President,

in the Chair.

THE Secrerary read the following report on the additions made
to the Society's Menagerie during the month of February,
1911 :—

The number of registered additions to the Society’s Menagerie
during the month of February last was 123. Of these 38 were
acquired by presentation, 47 by purchase, 8 were received on
deposit, 24 in exchange, and 6 were born in the Gardens.

The number of departures during the same period, by deaths
and removals, was 138.

Amongst the additions special attention may be directed to :—

2 Northern Lynxes (Felis lyna isabellinus), from Tibet, pre-
sented by Capt. D. G. Oliver, on Feb. 17th.

1 Dwarf Mongoose (Helogale varia), new to the Collection, from
Lamu, presented by the Rev. W. D. Braginton, on Feb. 27th.

1 Black-footed Polecat (Putorius nigripes), new to the Collection,
from N. America, received in exchange on Feb. 16th.

2 Cat-Bears or Pandas (4lurus fulgens), from Nepal, purchased
on Feb. 20th.

1 European Bison (ison bonasus), from Lithuania, presented

Proc. Zoou. Soc.—1911, No. XL. 40

558 MR. R. I. POCOCK ON

by H.G. The Duke of Bedford, K.G., President of the Society, on
Feb. 23rd.

2 Dybowski’s Deer (Cervus hortulorum), from Manchuria, new
to the Collection, presented by Sir Edmund Loder, Bart., F.Z.5.,
on Feb. 23rd.

1 Aldunati’s Finch (Phrygilus aldunatii), from Chili, new to the
Collection, presented by Miss Phillis True, on Feb. 13th.

1 King Penguin (Aptenodytes pennanti), from the Antarctic Seas,
presented by Sefior Clemente Onelli, on Feb. 11th.

Mr. D. Seru-Smrru, F.Z.S., the Society’s Curator of Birds, ex-
hibited a living Hybrid Duck, which was believed to be a cross
between the White-eyed Pochard (Aythya nyroca) and the
Marbled Duck (JZarmaronetta angustirostris). It was hatched at
Scampston Hall, Yorkshire, from a clutch of eggs laid by a Mar-
bled Duck at Lilford Hall, Northamptonshire. - The bird displayed
the characteristic markings of Marmaronetta, but the general
colour was dark reddish brown. It had been presented to the
Society by Mr. W. H. St. Quintin, F.Z.S.

Mr. R. I. Pocock, F.R.S., F.L.S., F.Z.8., Superintendent of
the Gardens, exhibited :—

(1) A pair of Otter cubs about seven weeks old, which were
found under a landing-stage at Tewkesbury, and were presented
to the Society by Mr. W. Baring Bingham, F.Z.S.

(2) The skin of an adult female Chacma Baboon (Papio porcarius)
representing an apparently undescribed local race of that species,
which he proposed to name and diagnose as follows :—

PAPro PORCARIUS, subsp. GRISEIPES *.
Abstract P. Z.S. 1911, p. 17 (March 28th).

Of the size, general appearance, and coloration of the typical
form from Cape Colony, but distinguished by the absence of black
hairs from the upper sides of the hands and feet, the extremities
of the limbs and tail being clothed with grizzled or annulated
hairs.

Loc. of type: Potchefstroom in the Transvaal.

This animal was presented to the Society in 1904 by Mr. James
Adams, whose son had brought it from the above-mentioned
locality. She was adult at the time and died in 1910. On
arrival she was seen to differ in the characters enumerated from
the ordinary South African Chacmas, commonly received by
the Society, many of which are shipped from Algoa Bay and all
of which are believed to come from some part of Cape Colony.
This belief is borne out by Mr. W. L. Sclater’s t description of

* The complete account of this new subspecies appears here, but the name and a

preliminary diagnosis were published in the ‘ Abstract,’ No. 93, 1911.—Ep1ror,
+ ‘The Fauna of South Africa: Mammals,’ i. p. 14, 1900,

A NEW CHACMA BABOON. 559

P. porcarius, in which it is stated that “the lower part of the
arms and legs |are| darker than the rest of the body, almost black,
the upper part of the hands and feet quite black.” Since
Mr. Selater further adds that “there are in the South African
Museum skins and skulls from the Stellenbosch, Tulbagh,
Worcester, Beaufort West, and Albany divisions of the Colony,”
it may be inferred that his description of the typical form of the
species was taken from the material in question.

During its lifetime in the Gardens, the baboon did not show
any change in colour, so that there are no grounds for believing
that the differences above pointed out are due in any way to age.
The Society also possessed at one time a second specimen, an adult
male, deposited by the Hon. Walter Rothschild, which exactly
resembled the type ; but for this, unfortunately, no precise locality
was known. The exact northern range in 8. Africa of the typical
or black-handed Chacma appears to be unknown; but on the
evidence supplied by the type specimen of P. porcarius griseipes,
it may be supposed that the latter supplants the former at least
in the Southern Transvaal. Chaemas have been recorded by
Mr. Vaughan Kirby fron the country between Beira and the
Zambesi; and according to Mr. Sclater the species is abundant
in Natal and the Transvaal; but it does not appear that any
specimens from the latter country have been described or critically
examined hitherto.

Text-fig. 145.

Black-footed Polecat (Putorius nigripes).

(3) A specimen of the North American Black-footed Polecat
(Putorius nigripes) (text-fig. 145), recently received in exchange
from the Zoological Society of Washington.

The peculiar coloration of this animal, taken in conjunction

40*

560 ON A BLACK FOOTED POLECAT.

with its behaviour in captivity, suggests that it belongs to one of
the warningly coloured species of Mustelide. Although the upper
surface of the head, neck, and body is tinted with buff, owing
to the terminal portion of the long hairs being that colour,
the whiteness of the underfur, and of the basal portion of the
long hairs of these areas, together with that of the sides and
under surface of the body and of the greater part of the tail and
face, gives the impression of a whitish animal which must be
conspicuous against any dark background. Sharply contrasted
with the creamy tint of the body is the jet-blackness of the legs
and of the tip of the tail; while the whiteness of the face is
emphasised by a broad black band stretching across the lower
part of the forehead and the base of the nose and involving the
bright bead-like eye on each side.

Apart from its coloration, this Polecat exhibits in un-
mistakable manner that fearlessness which is so marked a
feature of protected animals. It never makes any attempt to
keep in the background or lie hid, like a savage cat, when
disturbed, but comes boldly to the bars of the cage with threatening
aspect, as if eager to attack the intruder, uttering every now
and again a shrill chattering scream. Its obstinate pertinacity,
indeed, in keeping to the front when any human being is in
sight makes the opening and shutting of the door of the
cage for cleaning and feeding purposes a matter of serious
difficulty. Nothing but a bass-broom, which few small mammals
will face, has any effect in making it beat a retreat. Finally,
when actively interfered with by being netted, it emits the
foetid odour so characteristic of species of the genus Putorius.
The combination of characters here mentioned—namely, the
repulsive smell, the courage and fearlessness of exposure, the
conspicuous coloration—are all found in the Skunks, the stock
examples amongst the Mammalia of protected self-advertisers.
Added to these attention-arresting characters is the penetrating
scream comparable as a warning signal to the rattling of a
Porcupine’s tail. These facts justify the placing of the Black-
footed Polecat in the category of aposematic Mustelines, an
account of which has already been published by the Society *.

(4) The antlers of an old male of the Manchurian Wapiti (Cervus
aanthopygus), that recently died in the Gardens, to show the
natural variation in the position and growth of the second or
“bez” tine. In one antler this tine was much smaller than the
first or ‘‘ brow” tine, and was situated nearly midway between it
and the third or “trez” tine, apparently as in the specimen of
this Deer described by Mr. Lydekker as C. bedfordi; but in the
other antler the bez-tine was as long as the brow-tine and placed
close above it as in normal antlers of the stags of this group.

* R. 1. Pocock, P. Z. 8. 1909, pp. 944-959.

duth, sc. et imp.

IGS VOCs US WULOSWS Aerveleleveta,

deg 20S), IS}, IAL OVAGUL,

EW. Sextom del. Huth,sc.etmp.

1S So nh PE OCEANUS GUl TLGWS Grube On 22 UNG UlS es umpsions
LSS iG. ERS U LEMANS) Bate 7520. Sis Pin OS tis iNicwemlean,

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EW. Sexton, del,
LEPTOCHEIRUS PHOTINATUS Norman:

ON THE AMPHIPOD GENUS LEPTOCHEIRUS. 561

PAPERS.

26. On the Amphipod Genus Leptocheirus.
By E. W. Sexton, Marine Biological Laboratory, Plymouth *.

[Received December 22, 1910: Read March 21, 1911.]
(Plates XVIJ.-XIX.+, and Text-figure 146.)

The genus Leptocheirus was instituted by Zaddach in 1844, with
the type species Z. pilosus. So much discussion has arisen over
this species that it seemed desirable to settle the matter by refer-
ence to the actual specimens, should it prove possible to trace them.
Dr. Braun, in whose keeping at the Konigsberg Museum they were
discovered, most kindly sent them to me for examination and also
granted permission to dissect and figure one of the specimens, so
that the vexed question might be finally set at rest. JI am deeply
indebted to him, and to all those also who have so generously
assisted me in this investigation : to Canon Norman, for the loan
of specimens of ZL. subsalsus, pinguis, hirsutimanus, guttatus, and
pectinatus ; to Monsieur Chevreux for specimens of L. cornwauret,
tricristatus, and dellavallei ; to Dr. Kiikenthal and Dr. Zimmer for
permission to examine Grube’s specimens of Protomedeia hirsuti-
mana? and P. guttata; to Dr. Hjalmar Théel and Dr. Holmquist
for the trouble they have taken in searching for Ohlin’s specimen
ot LZ. aberrans ; to Professor Steuer for his assistance in tracing
Heller’s specimen of P. hirsutimana; to Dr. Otto Pesta for the
description and figures of this same specimen, by which its
identification became possible; and to Professor Vayssiere and
Monsieur Collin, of Cette, for the help given in searching for
Catta’s specimen of massiliensis.

It will be noted that the number of species has been reduced.

L. cornuauret Sowinski and subsalsus Norman become synonyms
of the type species, the one being the full-grown male, and the
other the full-grown female. Norman’s record is very interesting
with regard to the distribution, confirming Zaddach’s account,
both being for fresh or almost fresh water. All the other species
of the genus, so far as yet known, are marine.

I have also shown, I hope conelusively, the identity of
L. pectinatus Norman with L. dellavallei Stebbing, whick indeed
Mr. Walker has always maintained. JL. pectinatus is, in my
opinion, the young form sexually mature but not full-grown,
while Z. dellavallei is the fully developed animal. Figures are
given of the characters on which Chevreux (16) p. 91, and
Norman (36) pp. 87, 88, separated them, to prove that the
differences are merely those due to sex and development. The
question of distribution is also discussed by these authors in
separating the species, pectinatus having always been found near
the shore, and dellavallei at greater depths, but our present

* Communicated by Dr. W. T. Cauman, F.Z.S.
+ For explanation of the Plates see p. 593.

562 MRS. E. W. SEXTON ON THE

knowledge of the bathymetrical limits of any species is far too

inddequate to base any conclusions upon. Grube’s hirsutimana

from the Adriatic is the same species.

L. tricristatus Chevreux becomes a synonym of guttatus Grube.
An interesting point in this species is the varying development of
different characters, notably the antenne, the last perzeopod, and
the last uropods.

L. bispinosus Norman I consider identical with the species
described and figured by Della Valle as Z. guttatus. Heller’s
P. hirsutimana is to be referred to this species (see p. 585).

I have been unable to trace the type specimens of two species,
L. aberrans Ohlin, and Protomedeia hirsutimana var. massiliensis
Catta. The only specimen of aberrans, dissected by Dr. Ohlin for
the purpose of description, appears unfortunately to have been
lost.

There is nothing to add to Mr. Stebbing’s definition of the genus,
‘Das Tierreich,’ p. 625, except that the outer ramus of uropod 3 in
all the species is 2-jointed, the terminal joint rudimentary.

The development of the secondary sexual characters in the male
can only be definitely stated in two species, LZ. pilosus and pinguis ;
a great deal more material must be examined before it can be
decided whether the rule which applies to these species holds good
for the whole genus or not. In pilosws and pinguis the first gna-
thopod in the male is longer than the second, the hand is greatly
developed, the palmar margin concave; while in the female, on the
contrary, the first gnathopod is shorter than the second, and the
palmar margin is convex. The full-grown female, in all the other
species, agrees with this description. I find another distinction
between the sexes in the type species (Z. pilosws), in the shape
of the basal joint of the 5th pereopod (see Pl. XVII. figs. 22
& 23); but as this is the only species in which I have had the
5th pereopods of both sexes, I cannot say if this distinction is
generic or only specific.

The species now included in the genus are as follows, arranged
in chronological order, with their principal distinguishing cha-
acters :—

1. L. pilosus Zaddach 1844, = L. cornwauret Sowinski, 1898, the 3,
and L. subsalsus Norman, 1908, the 2. Pleon-segments not
dentate: accessory flagellum 1-jointed: gnathopod 1 greatly
developed in ¢, longer than gnath. 2; shorter than gnath. 2
in 2; dth joint in ¢ powerful and curved, 6th bent inwards
at right angles to the 5th, palmar margin concave ; palmar
margin convex in ?: finger of gnath. 2 almost straight, apex
acute.

2. L. pinguis Stimpson 1853. Pleon-segments 4, 5, & 6 each with
2 dorso-lateral angles : accessory flagellum long, 6—8-jointed :
hind margins of sideplates 1—4 serrate, spiniferous : hand of
gnathopod 1 greatly developed in ¢, and much _ longer
than gnath. 2; much shorter than gnath. 2 in 9 ; palmar
margin concave in d, convex in ?: finger of gnath. 2 as in
pilosus.

3 LD,

i=

L

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“ID

(

L.

Shp
i De

AMPHIPOD GENUS LEPTOCHEIRUS. 563

hirsutimanus Bate 1862 = Boeckia typica Malm 1871. Pleon-
segments not dentate, the 4th with a dorsal depression :
accessory flagellum 6- -jointed : sideplate 1 small, hidden by
the large sideplate 2: finger of gnath. 2 as in pilosus : fingers
of pereeopods 3-5 bifid : uropod 2 unusually massive.
guttatus Grube 1864 = Ptilocheirus tricristatus Chevreux
1886. Pleon-segment 4 with 3 medio-dorsal angles or teeth :
superior antenne not much longer than inferior ; accessory
flagellum 2-3-jointed ; palmar margin convex in 2: finger of
gnath. 2 unguiform, acute: falciform processes of uropods 1
and 2 of great length : inner ramus of uropod 3 tipped with
1 spine.
. pectinatus Norman 1869 = Protomedeia fasciata Costa 1864
and L. dellavallec Stebbing 1899: pleon-segments 4 and 5 each
with 2 dorso-lateral angles: sideplate 1 small, hidden by the
large sideplate 2: accessory flagellum 2—3-jointed : palmar
margin, gnath. 1, concave in ¢, convex in 2; finger of gnath.
1 exceeding the palm in length: finger of gnath. 2 straight,
laminar, tipped with setz.
aberrans Ohlin 1895.
bispinosus Norman 1908 = Protomedeia hirsutimana Heller
1866 and L. guttatus Della Valle 1893: resembles guttatus.
Pleon-segments 4 and 5 each with 2 dorso-lateral angles:
accessory flagellum 5-jointed: palmar margin, gnath. 1,
convex in 9: 2nd joint in gnath. 2 unusually long; finger
as in guttatus: falciform processes of uropods 1 and 2 of
great length: uropod 2 with two clusters of feathered sete on
the lower margin of the inner ramus.

Fam. PHorips.

For synonymy see Stebbing, Das Tierr. Lief. 21, p. 603.)

Gen. LEeprocHErRuS Zaddach.
(Stebbing, p. 625.)

LepTocHEIRus Prtosus Zaddach. (Plate X VII.)

1844.
1848.
1862.
1873.
1878.
1888.
1898.
1906.
1908.
1910.

Leptocheirus pilosus Zaddach (53) pp. 8 & 9.
wu Muller (32) p62:
Protomedeia pilosa Spence Bate (1) p. 168.
Leptochirus pilosus Mobius (80) p. 117.
Protomedeia pilosa Zaddach ( 54) 1 pp. 18 & 19.
Leptocheirus pilosus Stebbing (42) see p. 1707 for references.
45 cornuaureitSowinski(41) p. 470, pl. ix. figs. 9-22
3 pilosus Stebbing (44) p. 630.
é subsalsus Norman (38) p. 307, pl. xil. figs. 1-6.
Hf cornuauret = subsalsus Chevreux (18) p. 2.

This species was fully described by Zaddach in 1844, and the
accuracy of his description will be seen on comparing his account

with

the figures here given. In his later work, ‘Die Meeres-

564 MRS. E. W. SEXTON ON THE

Fauna an der preussischen Kiiste,’ is an interesting note on the
distribution of the species. It was found,so Zaddach states, both
in fresh and in salt water; by Rathke in 1843 in a large fresh-
water lake, the Geserich See, and by Zaddach himself, about the
same time, in the sea at Dantzig. It is not known under what
circumstances Rathke obtained his specimens, whether he collected
them himself near the banks, or whether they were given to him by
the fishermen. Zaddach, 34 years later, dredged the southern part
of the lake in search of the species, but his efforts proved fruitless.

The type * specimens are preserved in the Kénigsberg Museum,
and are labelled ‘‘ Protomedeia pilosa Zadd. Rauschen, Ostsee IX.
1866.”

The next record, also from the Baltic, is that of Miiller, who
noted the species as not rare in Greifswalder Bodden. He pointed
out the presence of a minute 1-jointed accessory flagellum on the
superior antenna, which had been overlooked by Zaddach.

The next authentic record of the species is in 1898, when
Sowinski described and figured the adult male as LZ. cornuwauret.

In 1908 Canon Norman described and figured the female under
the name of LZ. swbsalsus from specimens found by Mr. Gurney in
Norfolk, in fresh or almost fresh water.

In a note published in July, 1910, Monsieur Chevreux points
out that cornuaurei Sow. is the male of subsalsws Norman.

The examination of the type species which I have been enabled to
make and the comparison of it with specimens of both cornwawret
and subsalsws, prove beyond question the identity of all three forms,
pilosus being the young female, not sexually mature, cornuauwret
the full-grown male, and subsalsus the full-grown female.

The specimens examined were :—

Zaddach’s two types from the Baltic, measuring 4 mm.
around the curve; young females, not sexually mature.

4, 6 and 9, from muddy piles, R. Bure, Norfolk, Canon
Norman’s collection, measuring 5 mm. in a straight line.

4, $ and Q, from the coast of Algeria, sent by Monsieur
Chevreux, measuring 4—4°5 mm. in a straight line.

Description.

Body (Pl. XVII. fig. 1) smooth, moderately compressed ; pleon
without teeth, but last four segments with a seta inset on either
side of the median line.

Head about as long as the first two perzeon-segments ; lateral
corners rounded, not prominent.

Eyes oval in the small specimens, almost round in the large
ones. The ommatidia, which are large and few in number, are

* Prof. Braun informs me that these specimens were collected by Zaddach himself
in the Baltic at, Rauschen, about 35 km. northwest of Kénigsberg, in September 1866,
and are to be regarded as the types of the species. As the original description was
published in 1844, they are not, strictly speaking, types, but rather, perhaps, meta-
types.

AMPHIPOD GENUS LEPTOCHEIRUS. . 565

darkly pigmented in the centre of the eye; the outer ring in
the young animal is quite colourless, though this, of course, may
only be due to the fading of the pigment, little of which is left in
Zaddach’s specimens, while in Chevreux’s it is still fresh and
black.

Sideplate 1 free of sideplate 2, but not as deep and not quite
half its width ; obtusely rounded. Sideplate 2 the deepest of all ;
expanded inferiorly ; in Zaddach’s specimens and Norman’s small
one it is deeper than broad, but in the large specimens it 1s more
expanded in proportion to the other sideplates, in the largest of all,
a female, it is almost twice the width of sideplate 3; hind margin
straight, front angle produced and rounded. Sideplates 3 and 4
subequal, deeper than broad, of equal width throughout; front
and hind margins straight ; the 5th has the anterior lobe as wide
and as deep as sideplate 4, posterior lobe small; 6th and 7th
small ; 1-5 with a few piumose hairs and setules on the inferior
margin.

Pleon.—Segment 3 the largest; the 3rd—6th each with 2 sete,
one on either side of the median line, those of the 4th and 5th
segments the longest. These sete are inset submarginally on the
posterior margin in the same position as the angles or teeth of
the other species. Hind margin of the 2nd segment straight ;
that of the 3rd rounded ; inferior margins of 1-3 with plumose
sensory hairs, those of the 2nd longer and more numerous ; hind
margins 1-3 crenulate, 2-3 crenulations each with a setule inset.

ANTENNE (figs. 2 & 3).—Superior Antenna (fig. 2) much longer
than the inferior, about half the length of the body: 1st joint of
the peduncle stout; 2nd much more slender and longer than the
Ist in the large animal, in Zaddach’s smaller specimen it was
subequal to the Ist in length; 31rd not quite half the length of
the 2nd. Primary flagellum: Zaddach gives the number of
joints as 12, Norman as 13; one of Zaddach’s specimens, the
smaller of the two, had 11 joints, the other 12; Norman’s
largest specimen had 14; Chevreux’s 14, one specimen with 14
in one antenna and 15 in the other; the difference in number is
evidently due to development. The proximal joints are short,
the distal ones increasing in length, each, from the 5th or 6th to
the second last joint, being furnished with a stalked sensory
filament in addition to the small sete. Accessory flagellum not
as long as the Ist joint of the primary. It consists of 1 small
joint, which is so constricted near the apex as to give the appear-
ance of a minute second joint; furnished with 3 apical set, one
of great length, and 2 long setz above the constriction. Zaddach
completely overlooked this small appendage, which Muller was
the first to note, and indeed it might, as Miiller says, easily
escape attention, were it not for the long outstanding apical
bristles.

Inferior Antenna (fig. 3)—The proportions of the last three
joints of the peduncle are as given by Zaddach ; 3rd short, nearly
twice as broad as the succeeding joints; 4th the longest; 5th a

566 : MRS. E, W. SEXTON ON THE

little shorter than the 4th. The flagellwm is about half as long
again as the 5th joint; it is composed of 8 joints in Zaddach’s
specimens, of 10 in the larger ones, the terminal joint in all
microscopic.

OraL Parts.—Upper Lip (fig. 4): apex asymmetrically bilobed.

Lower Lip (fig. 5) large; outer lobes rounded, inner lobes
appear to be coalesced at the base.

Mandibles (fig. 6).—Both cutting-plates and accessory-plates
divided into 4-5 strong rounded teeth, the lowest being the
largest ; 7 dentate spines in the spine-row in Zaddach’s specimen,
9-10 in the large animals, the first two or three laminar. The
molar is large and cylindrical, crown ridged with rows of teeth,
and edged with flat spines, furnished with a long feathered seta
above ; in the figure the molar is not well represented, being
bent back to show the other portion of the mandible more clearly.
The palp is very large; 3rd joint the longest, tipped with 4
strong curved bristles, the distal inner margin furnished with a
double row of finely serrate bristles.

Masilla | (fig. 7) as described by Zaddach : inner plate large,
with 1 long plumose seta; outer plate with 11 strong spines on
its apex, 3 dentate ones at the upper angle, and the others
arranged in pairs, 1 bifurcate and 1 dentate together; in the
figure only one of the rows can be shown. The 2nd joint of the
palp widens towards the truncate apex, which is furnished with 4
strong short spines inset apically, and a diagonal submarginal
row of 4 sete.

Maxilla 2 (fig. 8) as large as maxilla 1; cnner plate the smaller ;
the apices of both plates carry long curved stiff bristles; the
inner margin of the inner plate has 2 rows of plumose sete,
setting out at different angles, one row containing 3 times as
many setz as the other.

Maxillipeds (fig. 9).—Inner and outer plates well developed ;
inner plates narrowed distally, the truncate apices inset with 3 flat
spines, outer margins with 3 setiform spines distally ; a row of 7
long jointed plumose setz crosses each of the plates diagonally
from the outer angle of the apex to the inner margin, extending
more than halfway down the latter. The outer plates are
widened distally ; in Zaddach’s specimen the plate on the right
side is furnished with 1 long plumose seta apically and 8 graduated
strong spines along the inner distal margin ; the plate on the left
has 2 of the plumose set apically, and 7 of the graduated spines ;
the outer surface carries numerous stiff curved bristles along the
inner margin. Palp, 2nd joint much the largest; 3rd much
produced on the inner surface over the insertion of the finger ;
finger small, obtuse, tipped with strong serrate setiform spines.
The 2 basal joints and the 1st-3rd joints of the palp are provided
on the outer surface with numerous long delicate plumose sete.

The Lirst Gnathopod (figs. 10-15) is longer than the second
in the male, shorter than the second in the female. It is
characterised by the remarkable development of the 5th—7th joints

AMPHIPOD GENUS LEPTOCHEIRUS. 567

in the male. In the female (fig. 11) the hand is shorter than the
preceding joint and lies in the same plane with it, but in the male
it is a quarter as long again as the 5th joint, and is carried bent
at right angles to it (fig. 12). The finger differs also: in the
male it is much stouter, more curved, and with a depression
in the inner margin proximally. The long 2nd joint is fur-
nished with a cluster of long delicate sete on the posterior
margin, and a row of short plumose setze on the anterior margin,
with another row of longer similar ones on the inner surface.
3rd joint bulging behind; 8rd and 4th fringed posteriorly with
long plumose sete ; 5th joint with transverse rows of plumose
sete, and 4 fan-like clusters of rigid serrate bristles on the
posterior margin. These bristles (fig. 15) are found on the 5th
joint of the first gnathopod in all the species of the genus. The
6th joint or hand is subequal to the 5th in length in the imma-
ture specimen (fig. 10); distally widened; palm slightly oblique ;
palmar margin microscopically pectinate, palmar angle with a
row of spines on the outer side, and 1 large spine on the under
side, with the tip of the finger fitting between them. The outer
row consists of 6 graduated slender spines, the first much the
Jongest, the shafts of which are produced apically into two unequal
processes, with a flat, delicate, feathered end-piece between the
processes. In one specimen there were 6 on one gnathopod and
7 on the other. On the inner side of the palmar angle a very
large stout sensory spine is inset, accompanied by 4 short stumpy
bristles similar in construction to those of the outer row. A row
of these stumpy bristles is found submarginally on the under
surface of the palm, and 3 or 4 rather larger ones on the outer
side. The finger in the young (fig. 10) is stout and curved, as
long as the palm, also microscopically pectinate, with a short
decurrent tooth near the apex, 2 sete inset in the notch, 1
smaller tooth behind, and 1 setule proximally. The full develop-
ment of the finger of the female is shown in fig. 13 and of the
male in fig. 14.

Second Gnathopod (figs. 16-18).—2nd joint very long, laminar,
with two rows of exceedingly long plumose set anteriorly,
one marginal, and the other submarginal on the under surface ;
posterior margin with only 3 or 4 long simple sete. A chitinous
ridge extends diagonally across the distal half of this joint, and
terminates at the posterior angle of the 3rd; this appears to be
the “crista” referred to by Zaddach (“in superficie externa
crista quadam a basi secundi articuli ascendente”). The 4th, 5th,
and 6th joints are practically subequal to each other in length ;
distal margin of the 4th joint with very long plumose sete on the
under side; posterior margins of the 5th and 6th with fan-like
clusters of short, stiff, finely serrated sete, anterior margins with
long plumose sete, the number of which increases a little with
growth, the smallest specimen having 9 on the 6th joint and
the largest 11, 3 on the 5th joint in all specimens. The
6th joint is slightly narrowed distally. The finger is almost

568 MRS. E. W. SEXTON ON THE

straight, the tip curved and 2 setules inset subapically. I have
given figures (see figs. 17 & 18) from both Zaddach’s and
Norman’s specimens drawn to the same magnification.

Pereopods 1 and 2 (figs 19 & 20) glandular; 4th joint very
much expanded, as noted by both Zaddach and Norman; 6th
joint as long as the 4th, but much narrower; 5th only half the
length; finger strong, two-thirds the length of the preceding
joint, with the gland-aperture at the tip.

Hinder Pereopods (figs. 21-23) rapidly increase in length; the
6th joint in all is much narrower and longer than the pre-
ceding. In pereopod 3 the joints are stout and short, furnished
with a few strong sensory spines ; 2nd joint obliquely oval, about
as wide as long, the anterior margin inset with 3 clusters of
jointed sensory ciliated hairs, posterior margin slightly crenulate,
with a setule in each crenulation, and with 4 plumose hairs
submarginally; finger about half the length of the 6th joint,
falciform. In pereopod 4 the 2nd joint is oval, longer than wide,
anterior margin with clusters of the sensory plumose hairs, pos-
terior Inargin crenulate, with a submarginal row of close-set long
ciliated hairs. In pereopod 5 there is a remarkable difference in
the shape of the basal joint in the male and female. In the
female the joint is oval, the posterior expansion narrowing distally,
while in the male it widens distally and is produced downwards
in a rounded lobe, but the curious part is that the ciliated sensory
hairs, instead of being inset close to the margin as in the female,
are at some distance from the margin on the under surface, but
yet give the same outline as in the female (cf. figs. 22 & 23).

Pleopods (fig. 24).—Inner rami twice the length of the peduncles;
outer rami the shorter, about three-quarters the length of the
inner. The peduncles of the 1st pair of pleopods carry each
on the outer side 9 long plumose hairs, on the inner side 1
exceedingly long plumose hair and 2 small coupling-spines with
recurved apices, the upper one (fig. 24) with 3 recurved teeth
on either side, and the lower with only 2. The peduncles of
the 2nd and 3rd pairs have only 1 or 2 long sete on the
outer side. The number of joints in the outer ramus is 11
in the Ist pleopods, 10 in the 2nd and 3rd; in the inner ramus
10 in the Ist, and 9 in the other pairs.

Uropods (fig. 25) extend backward to the same level; lst and
2nd pairs much alike in construction. The peduncle of the 1st
is as long as the inner ramus, with 3 or 4 spines on the
upper curve (3 in the small specimen, 4 in the larger), and
1 curved spine underneath, and the falciform apical process
reaching to half the length of the inner ramus ; outer ramus the
shorter, 1 spine inset on the upper margin, apex blunt with
a cluster of 4 stout spines; inner ramus with a similar apical
cluster and 2-3 along the upper margin. In uropod 2 the
peduncle is not as long as the inner ramus; it equals the
outer in length and carries 1 spine; rami as in uropod 1.
In uropod 3 the peduncle is subequal to the rami in length,
outer ramus, if anything, slightly longer than the inner, with a

AMPHIPOD GENUS LEPTOCHEIRUS. 569

cluster of setiform spines at the apex ; inner ramus with | spine
midway, and | spine and 2 setee at the apex; the large animals
have more spines ; all the spines are short and stout, “each with
an apical filament. The outer ramus is 2- “jointed, the terminal
joint rudimentary, carrying | long serrate spine or bristle.

Telson (figs. 25 & 26) short, broader than long, depressed
in the middle between the prominent lateral angles; apex
rounded ; 1 setaand 1 ciliated sensory hair on each angle, and
a pair of small ciliated hairs on either side.

Colour described by Zaddach as “ flavescens, dorso punctis
nigricantibus sparso.” In both his specimens and Norman’s the
colour has faded, but m Chevreux’s it is still fresh and vivid.
The whole animal is a beautiful golden yellow tint with stellate
markings in dark brown, These markings extend over the whole
dorsal surface of the head ; the anterior margin of the Ist, and the
posterior portions of the 3rd-7th pereon-segments and the Ist
pleon-segment are banded with them. The 5th and 6th pleon-
segments and the telson are entirely covered. An irregular band
of brown runs along each side of the perzeon just above the side-
plates and is continued along the epimera of the pleon to the
telson. All the side-plates and the posterior expansions of the
basal joints of the hinder pereeopods have each a patch of brown.

Distribution :—

GrsericH See, Prussia: and the Bauric: Zaddach (58)
I. pilosus, and (54) as Protomedeia pilosa.

Bauric : Greifswalder Bodden, Muller (32) as Z. pilosus.

Bauric: Mébius (30) as Leptochirus pilosus, depth 1-10 fins. ;
bottom, zostera and ulva.

R. Burn, Norfolk, England: Norman (38) as Z. swbsalsus.

The BospHorts: Sowinski (41) as LZ. cornuawrer.

Coast of Atgmria: Chevreux (18) as LZ. cornwaurei.

LEPTOCHEIRUS PINGUIS Stimpson. (Plate X VIII. figs. 10-12.)

For synonymy see Stebbing, Das Tierr. pp. 627 & 738, and
Norman (38) p. 309.

There is little to add to the summary of characters given by
Stebbing. The few notes subjoied were taken from three slides
kindly lent: to me by Canon Norman, and prepared by him from
specimens from Vineyard Sound and Long Island, N. America.

Head, 9 , lateral corners not much produced, truncate.

Superior Antenna.—I|st and 2nd joints in ¢ and @ practically
subequal to each other in length; 3rd a little move than one-
third as long as 2nd. Primary flagellum, 2, broken, 21 joints
remaining, all, except the first 6, with a small sensory filament
and short sete; 6 very long joints in the accessory flagellum which
equals 3 joints of the primary in length. In the male, the primary
flagellum is composed of 31 joints, Gah except the first 5, with
a small sensory filament. The accessory is broken, 6 joints
remaining, equalling 5 joints of the primary in length.

570 MRS. E. W. SEXTON ON THE

Inferior Antenna, 9 .—8srd joint short and broad, with 2 sensory
spines on the upper distal angle; 4th joint the longest; flagellum
longer than 5th joint, equal to 4th ; 11-jointed in this specimen,
each joint with a cluster on either distal angle of setiform spines
and very long set.

Lower Lip, 2, outer lobes rounded, densely setose ; inner lobes
coalesced, large, distally narrowed.

Mandible, 2 : cutting-plate on the right side produced below,
margin divided into 4 rounded teeth, the lowest the largest ;
accessory-plate much as in pectinatus, tapering from the base to
the long acute tip, with 2 very minute teeth above; 17 spines in
the spine-row, the first 4 large, the last 2 very small. Molar
prominent ; crown reniform in shape, edged with strong teeth,
and carrying a small accessory process on the side nearest
the cutting-plate. Palp: 2nd joint longer than Ist; 3rd
the longest, curved, falciform, attenuated distally, and armed as
in the type species with 4 long stiff curved bristles apically,
2 dense rows of similar shorter bristles on the inner margin,
and about 4 clusters on the outer margin. The left mandible
was broken.

Mazxilla, 1 2, much as in the type species; outer plate in one
maxilla with 11 strong spines, 3 dentate ones at the upper angle,
and the others set in pairs, 1 bifureate and 1 dentate together ;
in the other maxilla there were 12 spines.

Mazwilia,2 2, as in pilosus; inner plate narrowed distally.

Mavillipeds, 2 , covered with numerous long set. The basal
joints, and the Ist joint of the palp, on the outer side carry
fan-shaped groups of exceedingly long plumose sete; the outer
side of the 2nd joint of the palp is also covered with them, and
carries in addition a fringe along the inner margin. Outer plate
expanded distally, densely fringed on the inner margin with
slender spines, the apical ones of great length. Jnner plate
elongate, tapering distally to the narrow truncate apex; 3 flat
spines are inset in the apex, almost hidden by the plumose setz
surrounding them; the outer margin is furnished with a row of
about 10 simple setiform spines, and the inner carries a row of
18-20 very long, flexible, plumose setze. The 2nd joint of the
palp wuch the longest, much wider and about twice as long as
the 3rd; the 3rd produced over the 4th as in the type species ;
4th small, tipped with strong serrate spines,

First Gnathopod, 9 .—5th joint very long, provided on the
posterior margin with transverse rows of feathered sete, and
the clusters of stiff serrate bristles found in all the species of
the genus; 6th joint subequal to the 5th in length, as described
by Stebbing ; palmar margin convex, serrate. On the outer side,
extending from the angle nearly to the middle of the palm, is a
graduated row of bristles, 7 in number, similar in structure to
those of the type species; a row of smaller bristles is inset
submarginally along the palm. On the inner side, at the palmar
angle, is a short, very broad, sensory spine, and a submarginal

AMPHIPOD GENUS LEPTOCHEIRUS, 571

row of about 17 very small bristles, like those of the outer side.
The finger is exactly the length of the palm ; being closed in the
specimen examined, the detail could not be seen, owing to the
palmar spines obscuring it.

This gnathopod in the male differs from that of the female, as
in the type species. The 5th joint is longer in proportion ; the
2nd and 6th joints shorter. The 5th joint equals the basal joint
in length. The 6th is not quite two-thirds its length, and in the
natural position is held bent almost to a right angle with the
preceding joint. It is expanded distally; front margin very
thick; hind margin laminar, convex on the upper surface, and
furnished with six transverse rows of sensory sete ; palmar
margin transverse, concave instead of convex, furnished with the
submarginal rows of bristles along the margin, and the graduated
group at the angle as described for the female; the very stout
strong spine defining the angle is much larger than in the female.
On the under side of the hind margin of the 5th joint (not the
“basal joint” as given by Holmes (26) p. 522), distally, a chitinous
spine-like process is developed; another similar but smaller one is
on the 6th joint proximally, with a groove beside it, into which
the larger one appears to fit when the hand is bent in. In order
to show these processes, the hand is represented (fig. 10) with the
hind margin uppermost, and, in consequence of being placed in
this position, the true measurements of this joint cannot be seen ;
viewed from the side, it is exactly the same width proximally as
the 5th, expanding gradually to the palmar margin. The finger
in the male i is much more arched than in the female: and w fea
closed, the tip, instead of meeting the angle as in the female,
impinges against the under surface of the hand; it is finely
serrate, with a row of setules inset, and apparently a small
auxiliary tooth near the apex, but this was too obscured hy the
overlying palmar bristles to be ascertained with any certainty.

Second Gnathopod (fig. 11) as figured by Norman. The finger
is about half as long as the preceding joint, lightly curved,
acute, of the same structure as in the type species; with 4 sete
on the inner margin, and a cluster of 3 subapically in the male ;
fewer setee in the female.

Pereopod 5.—The finger is of unusual length, considerably
longer than those of perzopods 1 and 2; very slender, alinost
straight, with a setule near the apex.

Or -opod 3 (fig. 12), 9 A figure is given of the rudimentary
2nd joint of the outer ramus, with the spines omitted.

LEPTOCHEIRUS HIRSUTIMANUS Spence Bate. (Plate XVIII.
figs. 13-16.)

The synonymy as given by Stebbing, Das Tierr, p. 627, omitting
the references to Heller 1866, and Grube 1866, and adding : =
L. pilosus Norman and Scott (36) p.

The animal described by Heller as Bie Soe | hirsutimana is

572, MRS. E. W. SEXTON ON THE -

a female of Z. bispinosus (p. 585): Grube’s specimens proved on
examination to be pectinatus (see p. 577).

Sars has given an excellent description and figures of this
species, (40) pl. 197, to which it is only necessary to add one or
two details.

Superior Antenna, flagellum furnished with very long sensory
filaments.

First Gnathopod (fig. 13).—The palm is oblique, serrate, with
its limit defined, as in pectinatus, by a large sensory spine inset
on the under side. The tip of the finger reaches to this spine.
The palm carries a small strong spine on the outer side, close to
the insertion of the finger.

Second Gnathopod.— The finger (fig. 14) is exactly as in the
type species, cf. Pl. XVII. figs. 17 & 18.

Hinder Perceopods.—Sars figures the finger of perzeopod 5 only
as bidentate, but the finger in all three hinder pereopods is of
the same structure (see fig. 15). Spence Bate in his original
description notes it for the 3rd perzeopod, (1) p. 169; and Malm,
(27) p. 546, describes and figures it for all three.

Uropod 3 (fig. 16).—The outer ramus is 2-jointed, as in all the
other species of the genus, but the 2nd joint is exceedingly small
and very difficult to observe except from the dorsal view, owing
to the apex of the Ist joint being produced beneath it.

LEprocHEIRus Gurratus Grube. (Plate XVIII. figs. 1-9.)
1864. Protomedeia guttata Grube (23) p. 63.

1866. ms - », (24) p. 408, pl. x. fig. 3

1885. si Camus: (6) go 7.

1886. Prlnchere us tricristatus Chevreux (9) p. xl.

1887. ™" i LO) ies

1887. a oa (11)p. 310, pl.v. figs. 3 & 4,

and fig. 4 in the text, p. 578.
1888. Leptocheirus guttatus Stebbing (42) p. 366.

1898. 7. 7 Ghevreux (15) p. a
1900. he (16) p.
1906. 53 guitatus Stebbing (44) p. oh
tricristatus Stebbing (44) p. 629.
1906. = guttatus Norman & Seote (36) p. 85, pl. ix.
figs. 4-7.
1907. 5 guttatus Norman (87) p. 369.
1910. e tricristatus Chevreux (18) p. 2

The specimens examined were :—

1 Q, Grube’s type specimen, 5°5 mm. long, from Breslau
Museum.

3 2 L. trieristatus, taken by Monsieur Chevreux in the Bay of
Quiberon, the largest measuring 4°5 mm.

9 taken by Canon Norman at Falmouth, the largest 9?
measuring 6°25 mm.

The type specimen from Breslau has Dr, Grube’s original label

AMPHIPOD GENUS LEPTOCHEIRUS. 573

still on the bottle—‘* Protomedeia guttata Gr. m. Hiern, Lorenz,
Val Cassione 36.” It is a female, 55mm. in length from the
tip of the rostrum to the tip of the telson, with the pigmentation
still vivid after 48 years in alcohol. I have described this
specimen in detail, the account of the specific characters given by
Grube not being sufliciently adequate for the present system of
classification. In the figure of the whole animal it must be noted
that as all the appendages were drawn in situ, the ewact
measurements and proportions of the joints of the pereopods
cannot be expected. Hach character is compared with Chevreux’s
specimens of tricristatus to prove the identity of the two forms.

Description.

Body (Pl. XVIII. fig. 1) more stoutly built than in pilosus and
pectinatus, and sideplates shorter in proportion than in those
species.

Head longer than the Ist and 2nd pereon-segments taken
together ; lateral corners not prominent, rounded.

Hyes round, black or brownish-black in colour ; ommatidia large.

Sideplates 1-5 subequal to each other in length. Sideplate
1 not covered by sideplate 2; produced forward over the side of
the head, of equal width throughout ; inferior margin obtusely
rounded, with 3 or 4 stiff, sparsely feathered sets inset; hind
margin, as in the type species, not continuous proximally. Side-
plate 2 the largest, a little expanded distally, front angle and
inferior margin rounded, hind margin straight; inferior margin
fringed with a row of flexible, finely plumose sete. Sideplates
3 and 4 alike in form, deeper than broad, slightly wider proximally ;
sideplate 5 with the anterior lobe about as deep as preceding
sideplate, but not as wide; inferior margins of all three carry a few
setze, similar to those on sideplate 1. Sideplates 4 and 5 small.

Pleon.—Segment 3 much the largest, as long as segments | and
2 taken together. The 4th segment has the armature characteristic
of this species; the posterior margin is produced in 3 processes,
the median one upstanding, acute, shorter than the lateral ones;
the latter broad at the base, much produced over the succeeding
segments, apices acute, curving upwards, with a spinule inset in
each. Segments 5 and 6 are small, the posterior margin of the
5th with a few setules dorsally. The hind margin of the 2nd
epimeron is straight, with 1 crenulation and setule at the postero-
lateral angle; inferior margin densely fringed with long plumose
sete. The hind margin of the 3rd is produced and rounded, with
2 indentations, a setule in each; inferior margin lightly concave,
with 3 long plumose sete inset midway, and 3 short spines near
the antero-lateral angle.

Antenna. Superior Antenna (figs. 2-4).—1st joint of the peduncle
stout, shorterthan the 2nd; 3rd not quite half the length of the 2nd.
Primary flagellum 7—-12-jointed, the largest female had 11 joints ;
accessory flagellum 2—3-jointed. I must note here an occasional

Proc. Zoot. Soc.—1911, No. X LI. Al

574 MRS. E. W. SEXTON ON THE

curious variation in the number and proportions of the joints of
the flagella in the same animal. This variation occurs, I believe,
not infrequently in the Amphipoda, some species of Jassa, for
example, having usually one joint more on one side than on the
other. In Grube’s type specimen the primary flagellum on the
right side has 8 joints, 7 on the left; the right flagellum of the
inferior antenne has 4, while the left has only 3; the accessory
flagella are both 2-jointed, but the one on the left antenna is
longer than that on the right. Grube evidently counted the
joints of the antenne on the left side. In two other specimens
examined, females of the same size, the superior antennz were the
same length, but in one case there were 12 joints, and in the other
only 9; the accessory flagellum in both was 3-jointed, but whereas
in the animal with 12 joints it exceeded the 2nd of the primary
in length (fig. 4), in the other it did not reach to the level of the
2nd. Figures 2 and 3 are from another specimen, the right
accessory of which had 2 joints, the left 3. All the joints of the
primary, except the first 4, are provided with a sensory filament
in addition to the sete.

Inferior Antenna.—A4th joint of the peduncle the longest ;
flagellum a little longer than the 5th, subequal to the 4th in length.
Flagellum 3-6-jointed, the first the longest.

First Gnathopod (fig. 5).—Hand about twice as long as broad ;
palm oblique, defined, as in pectenatus, by a large sensory spine
inset on the under surface ; palmar margin convex, serrate, the
serrations turning the corner and ending at the level of the spine,
it carries 5 or 6 small sharp spines subinarginally on the outer
surface, and 8 smaller ones on the inner, all microscopically
serrate. The finger is as long as the palm, curved, serrate, with
2 auxiliary teeth.

Second Gnathopod (fig. 6).—2nd joint very long, equalling in
length joints 4-6 taken together. The 4th joint, as is usual in the
genus, is much produced over the 5th on the inner side, and if the
measurements are taken along this side, the 4th, 5th, and 6th are
seen to be subequal to each other ; Chevreux’s figure (pl. v. fig. 4
shows the outer side. The finger is about three-fifths the length
of the preceding joint, lightly curved, with 2 setules near the acute
apex.

MESReonae 1 and 2 alike in structure, glandular, resembling the
type species in the shape and proportions of the joints except that
the 5th joint is slightly longer in proportion than in pilosus, and
the finger is much longer and more slender, being equal in length

to the posterior margin of the preceding joint.

Hinder Per manos very like those of the preceding species, but
stouter and much less elongate. Norman gives ‘‘the greatly pro-
duced last pereeopod” as a specific character, but in most of the
animals examined by me the proportions are much as figured in
Grube’s type specimen (fig. 1); in the others the 6th joint is more
developed. The basal joint in all is large, rounded oval, produced
downwards in a lobe, furnished anteriorly with short spines at
intervals and ciliated tele s,and adense cluster of the latter at the

AMPHIPOD GENUS LEPTOCHEIRUS. 575

distal angle; the posterior margins of the 3rd and 4th joints are
crenulate, 6-9 crenulations each with a setule. In the 5th this
margin has 5-7 serrations, and 1-3 ciliated hairs inset in the
lobe. The 3rd joint carries a dense cluster of ciliated hairs at
the anterior distal angle; and the 5th joint a similar cluster
of feathered sete. Fingers short and curved. In peropod 3
the 4th joint is slightly longer than the 6th and wider;
5th shorter than 4th; 6th with groups of short spines. In
pereopod 4, the 4th and 6th joints are subequal to each other
in length; in one or two of the larger specimens the 6th is
slightly the longer; it carries numerous spines and a cluster of
very long setz at the distal angle. In perzeopod 5, the 4th, 5th,
and 6th joints rapidly increase in length; the 6th is slender and
lightly curved, inset on its posterior margin with exceedingly long
setze in addition to the spines.

Pleopods much as in pectinatus, except that the peduncles are
shorter in proportion to the rami, about one-third as long, and the
outer ramus is only two-thirds the length of the inner ; cleft spines
asin pectinatus, 5 in the 2nd pleopod; coupling-spines with 2 rows
of recurved teeth in addition to the recurved apex, 3 teeth in a row
in the lower spine, 2 in the upper.

Uropods (figs. 7 & 8).—There is a marked variation in the
length of the uropods in different specimens, most noticeable in the
3rd pair, connected apparently with the variation in the develop-
ment of the last pereeopod. In Grube’s specimen (fig. 8) the
peduncle of uropod 3 is short and broad, shorter than the outer
ramus; inner ramus not quite half the length of the outer. Some
of Norman’s and Chevreux’s specimens are like Grube’s type; in
others, the peduncle equals the outer ramus in length and the
inner ramus is nearly as long as the outer; while in the largest
specimens with the last perzeopods much produced, the peduncle
of uropod 3 is longer than the peduncle of uropod 2, and longer
also than the rami, both rami long and slender, inner quite equal
to the outer in length. Uropod 1, peduncle shorter than outer
ramus, the falciform process of unusual length, reaching almost to
the tip of the outer ramus, furnished with 5 spines along the
upper curve; outer ramus a little shorter than the inner, with
3 spines and an apical cluster of 5, 3 of which are longer than
the others; inner ramus with 4 spines and a similar apical group ;
these spines are longer and more slender than those of uropod 2.
Uropod 2 stoutly built, peduncle short and stout, with the falci-
form process equalling the outer ramus in length; outer ramus
shorter than the inner, with 2 spines, inner with 3, each with an
apical cluster of 5 strong stout spines (fig. 7). Uropod 3,
peduncle produced underneath in a flat laminar expansion; outer
ramus with a rudimentary 2nd joint carrying | long stiff sparsely
feathered bristle; the lst joint has a group of graduated similar
bristles almost concealing the terminal joint; inner ramusslender,
tipped with 1 stout spine. This description of uropod 3 applies
to all the specimens examined by me,

Telson (fig. 9) as figured by Chevreux and Norman: the apical

4\*

576 MRS. E. W. SEXTON ON THE

margin is convex; from the dorsal view it appears truncate,
cf. also bispinosus p. 089.
Colowr.—Chevreux in his original description gives the colour
as yellowish with some brown spots; in his later account he
describes it as generally uniform yellow, some specimens with
brown bands dorsally. In Grube’s type the colour is still vivid,
tawny yellow, with round stellate markings in light and fins
reddish-brown. These markings occur on the head ; in transverse
bands on segments 2-10; on the sideplates ; and on the posterior
expansion of the basal joints of the hinder peropods.
Distribution :—
CuanneL Isuanps: Norman (87) p. 369, as LZ. guttatus.
Fatmoutu Harsour: Norman & Scott (36) p. 86, as L. guttatus.
Oceanic Coast of France: Chevreux (11) p. 311,as Ptilocheirus
tricristatus, depth 7 m.; bottom deposit, gravel with coral-
lines; (12) p. 578, depth 10 m., bottom deposit, nullipores ;
(15) p. 481, as Z. guttatus, depth 10-50 m.

MEDITERRANEAN :—~ Coast of France: Chevreux (16) p. 92, as
L. guttatus. Coasts of Augmrta & Tunis : Chevreux (16) p. 92,

as L. guttatus; (18) p. 3. Apvriatic: Grube (24) p. 408, as
Protomedeia guttata.

LeprocHEIRus PecTINATUS Norman. (Plate XTX.)
The principal references to this species are as follows :—
1864. Protomedeia fasciata (non Kroyer 1842), Costa (19) p. 155,
pl. 1. fig. 8.
1864. Protomedeia Kr.
1864. Leptocheirus pilosus Grube (22) p. 73.
1866. Protomedeia hirsutimana? Grube (24) p. 402.
_ pilosa Grube (24) p. 417, pl. x. fig. 2.
1869, 3 pectinata Norman (34) p. 283.
1887. Ptilocheirus pectinatus Chevreux (11) p. 309.
1888. Leptocheirus pectinatus Stebbing (42) p. 1707 for references.

1893. A pilosus Della Valle (21) pp. 427-430, pl. iv.
fig. 10; pl. xii. figs. 1-14
1895. ss ,, Walker (47) p. 470.
1895. » (48) p. 310.
1899. f dellavallei. Stebbing (48) p. 350.
1900. 24 pilosus Chevreux (16) p. 90.
3 fasciatus Chevreux (16) p. 91.
1906. Be dellavallei Stebbing (44) p. 628.
i. pectinatus Stebbing (44) p. 629.
1906. . . Norman & Scott (36) p. 87, pl. ix.
figs. 1-3.
4 fasciatus Norman & Scott (36) p. 88, pl. v.
figs. 11 & 12.
1908. is dellavallei Norman (38) p. 310.

Bt pectinatus Norman (38) p. 310.

1909. x pilosus Walker (51) p. 341,

AMPHIPOD GENUS LEPTOCHEIRUS. 577

The specimens examined were :—

3 L. pectinatus, two females and one male, measuring 2°5—3 mm. ;
from Guernsey, sent by Canon Norman.

1 specimen, 9 , 4:°5 mm., taken by the ‘ Huxley’ in 109 fathoms,
during her cruise on the north side of the Bay of Biscay, 1906.

3 L. dellavallei, §, 5-6 mm., from the coast of Senegal, from
Monsieur Chevreux.

3 specimens from Breslau University Museum, referred to
above. These latter specimens were sent in two bottles with
Dr. Grube’s original labelsstill on them. One bottle, marked
““ Protomedeia pilosa Zadd. 9 . Luss. pice. Mundtheile. Grube,”
contained two tubes, with one specimen in each. One of these,
a female almost ready to moult, with a brood of young just
hatched, was dissected, and is evidently the specimen from
which Grube made his drawings. That this is so can be
proved by a comparison of the figures with the dissections:
e.g., the shape of the 2nd maxilla in his figure is due to the
fact that a portion of the inner plate with most of the
feathered bristles had been torn away (cf. (24) pl. x. fig. 2 m’,
with fig. 10); and again, in the Ist maxilla he notes the.
absence of the apical seta, which seta is, however, there, but
too completely masked by dirt to be seen, except under a high
power. The other tube contained a female 5°5 mm. long.
The second bottle was originally marked ‘“ Protomedeia
hirsutimana Sp. B. 2. Vollst. Luss. pice. Grube,” but over
the “hirsutimana ” is written in a different ink “ pilosa Zadd.”
and the words “‘m. Hiern ” added, apparently by Grube
himself. The tube in this bottle contains a large, brightly-
coloured female, the largest specimen I have yet seen.

The specimens form a most interesting developmental series 1n
the order in which I have arranged them above. Figures 14, 19,
& 23 are taken from a female pectinatus 2°75 mm., figs. 1 &
13 from a male 2°0 mm. long, Norman’s specimens: figs. 5, 16.
17, 25, & 28 are from the female specimen described by Grube:
the other drawings are from two males, dellavallei, 5°25 and
6 mm. respectively, Chevreux’s specimens. The pectinatus figures
are more magnified than the others for the purpose of comparison.

Description.

Head not quite so long as pereon-segments 1 and 2 taken
together ; lateral corners not prominent, truncate.

Eyes almost round, a little drawn out towards the lateral angle;
ommatidia few in number, large, with blackish-brown pigment
in the centre, outer row not so darkly pigmented.

Sideplates (figs. 13-16).—The first sideplate, which has been
the principal character for separating the two forms pectinatus
and dellavallei, is of exactly the same structure in all the
specimens. It is small, and completely hidden by sideplate 2.

578 MRS. E. W. SEXTON ON THE

(Through some mischance, Norman’s figure of the first gnathopod
((36) pl. ix. fig. 2) shows the second sideplate attached instead of the
first.) The first sideplate is subquadrate in the young animal, with
1 long sensory spine inset at the anterior angle, but with growth
this angle becomes more produced downwards, until in the largest
specimen of all the anterior margin of the sideplate is half as long
again as the hind margin. Four stages of development are
represented in the figures. The ‘Huxley’ specimen forms the
link between figs. 14 & 15; the anterior angle is not so much pro-
duced as in the latter, 6 setules are inset anteriorly, 1 inferiorly,
and 3 short sensory hairs at the posterior angle. In Grube’s
specimens, the anterior margin is lightly concave and the posterior
angle has a strong chitinous margin. The hinder portion of the
sideplate is firmly affixed to the basal joint beneath, so firmly in
fact as to make it impossible to separate them without destroying
the shape of the sideplate. A delicate laminar plate extends
beyond the sideplate behind, and is all but continuous with it
proximally, the line of demarcation being barely distinguishable.
Sideplate 2 is the largest of all, as deep as broad in the small
specimens, a little deeper than broad in the fully developed
‘ animal; front margin rounded; hind margin almost straight with
4 small sete inset; inferior margin rounded, thickly fringed
with delicate, sensory, cleft-tipped sete of varying lengths
(26 in the young, to about 41 in the full-grown). The remaining
sideplates are very like those of the type species. Sideplates 3
and 4 are subequal to each other in length, shorter. than the 2nd,
about half as deep again as broad. The 3rd is of equal width
throughout, front and hind margins straight, the latter with three
or four sete inset; inferior margin convex, fringed with 10-19 of
the sensory cleft sete. The 4th is a little wider proximally than
the 3rd, and the front margin is lightly convex; inferior margin
with fewer sensory sete, 5-15. In sideplate 5 the anterior lobe is
about as broad and as deep as the preceding sideplate; inferior
margin rounded, with only 4-7 sete; posterior lobe very small,
only one-third the depth of the anterior, 2 small sete inset.
Sideplate 6 small; posterior lobe about half as deep as the
anterior, with 1 small seta behind and 1 of the sensory
serrate spines similar to those found on the hinder perseopods
and uropods (see fig. 21); on the anterior lobe are 2 ciliated
hairs. Sideplate 7 small and subquadrangular.

Pleon._Segment 3 much the longest, as long as the 1st and 2nd
taken together; 4th segment a little depressed dorsally. The
hind margin of the 2nd is straight, that of the 3rd rounded;
inferior margins of 1-3 thickly beset with long plumose sete,
most numerous on the 2nd. On either side of the median line of
the 4th and 5th segments the posterior margin is produced in an
erect membranaceous lappet-like process, too delicate in structure
to be termed a tooth, each process having a setule inset in the
notch (fig. 26). Both Costa and Grube refer to these setules,

AMPHIPOD GENUS LEPTOCHEIRUS. 579

without apparently noticing the processes*, but this oversight is
very easily understood. The integument is so thin that if by any
chance, such as pressure or a little mucous dirt collected, the
process be flattened against the body, it is impossible to see it.
The 4th segment has a group of spines on either side, just above
the insertion of the uropods. The processes on the 5th segment
extend beyond the very small 6th segment.

ANTENNA. Superior Antenna (figs. 1—3).—1st joint of the peduncle
stout, shghtly longer than the 2nd in actual measurement, the
greater apparent length of the latter being due to its slenderness ;
in only one of the specimens, Grube’s largest, the 2nd joint
was a very little longer than the Ist; on the outer side are
several small ciliated hairs, and a cluster of sete with 1 long
ciliated hair at the distal angle; the inner angle carries a very
long, stout, outstanding sensory spine with 2 smaller ones inset
beside it. The 2nd joint is only half as broad as the Ist, with a
cluster of 3 or 4 sete and 1 long ciliated hair on either
distal angle. The 3rd joint in all the large specimens I have
examined barely reaches half the length of the 2nd; in the
smallest one it slightly exceeds half the length. The primary
flagellum is composed of 9-14 joints ; one young ovigerous female
2°5 mm. long had 9 joints; a male measuring 6 mm. had 14;
Grube’s large females 5°5 mm. and 6°25 mm. had respectively 12
and 10, the joints in the last-mentioned case being longer than in
the other animals examined. Each joint, from the 4th in the full-
grown and from the 2nd in the young, to the penultimate, is
provided with a very long sensory filament in addition to the
small sete. The accessory flagellum is usually 2-jointed, equalling
the Ist jot of the primary in length, and is so described by
Norman, Della Valle,and Walker. All the specimens examined
by me, except one, agree in having 2 joints only, though the
length varies a little, in the small specimen (fig. 1) being slightly
less than the Ist joint of the primary in length, and in the
medium-sized specimen (fig. 2) slightly more. The exception is
Grube’s largest specimen, which has 10 joints in the primary, and
a 3-jointed accessory flagellum equalling the first 2 joints of the
primary in length. Costa’s observation agrees with this, (19)
p- 155, “Il filetto composto di dieci articoli finamente pelacciuti ;
il filetto accessorio lungo appena quanto due articoli del primario.”
Grube in his description, (24) p. 403, says: ‘‘ Bei dem von mir
zuerst untersuchten Exemplar fehlte den oberen Antennen die
Nebengeissel..... An einem zweiten Exemplar fand ich die
Nebengeissel und zwar eine 3-gliederige.” The first specimen
examined by him (from Lussin-piccolo) had, however, an accessory
flagellum, a 2-jointed one, but this, in both antenne, had lost the

* “Quelli del quarto [abdominal segment] in oltre guerniti di piccole spine lungo
il margine dorsale.” Costa (19) p. 15.

“* Auf dem Riicken des 11ten und 12ten Segments vor dem Hinterrande sieht man
ein paar Borsten.” Grube (24) p. 407.

580 MRS. E. W. SEXTON ON THE

long apical setz, and, lying flat against the primary, was hidden
by a coating of dirt. In his third specimen both the superior
antennz are missing.

Inferior Antenna.—3drd joint of the peduncle short, as broad as
long, with a cluster of very long sete on the inner angle and 1
long sensory spine above. The 4th joint is twice the length of
the 3rd, slightly longer than the 5th, both beset with clusters of
the long, cleft-tipped setze and smaller setiform spines. Flagellum
shorter than the 5th joint, composed of 4 joints, the 1st much the
longest, almost equalling in length the two following taken
together, each carrying, in addition to the sete, a pair of strong
spines setting out on either side. In the small specimens the
flagellum is 3-jointed.

OrAL Parts.—The description is taken from the fully adult
specimens 5-6 mm. long, but the structure is precisely the same
in the small animals, the only difference being the lesser number
of spines and sete.

Upper Lip (fig. 4) thick, subquadrate in form, much arched
above; apex emarginate, slightly asymmetrical. In the figure the
lip is turned a little upwards to show the emargination of the
apex.

Lower Lip (fig. 5) as in the type species.

Mandibles (fig. 6).— Cutting-plate on the right mandible curved,
margin divided into 3 teeth, the lowest much produced; in
two specimens the margin was entire, the middle tooth not being
developed; accessory-plate narrow, in some specimens scarcely
wider than, and not as long as, the Ist spine of the spine row,
produced below to an acute tooth covered with microscopic
tubercles, and provided with a small tooth above. In the left
mandible the cutting-plate has 4 teeth, the second small, the
lowest the largest; the accessory-plate is strong, much broader
than that of the right mandible, the mandible divided into
3 teeth, the upper and lower of which are large, the middle one
small. There are 7 spines in the spine-row in Chevreux’s
specimens, 8 in Grube’s largest, the first 3 being unusually
large, wide at the base, flat and furry in appearance owing to
being covered with microscopic spinules. The molar is very
prominent; the crown transversely ridged with rows of teeth,
edged with flat spinules, with 1 long, feathered seta above.
The palp is very large; 3rd joint the longest, tipped with long
serrate bristles, with a double row of smaller ones extending
down the distal half of the inner margin; the outer margin has
4 groups of long serrate bristles inset.

Mazxilla | (figs. 7-9).—Jnner plate large, with 1 long plumose
seta inset; Grube notes the absence of this seta (p. 404),
but his specimen is so covered with mucous dirt as to render it
difficult to see details clearly ; not only is the seta present, but the
new one can be seen under the old loose skin, the animal, as
before stated, being ready to moult. Outer plate curved, with
11 strong spines, 4 of which are bifurcate (fig. 8), the rest

AMPHIPOD GENUS LEPTOCHEIRUS. 581

finely dentate (fig. 9); the outer margin is covered with fine
hairs. The 2nd joint of the palp is expanded apically, and
rounded, with a marginal row of 4 strong flat spines, and 3
plumose sete; 2 or 3 diagonal rows of set are inset sub-
marginally.

Maxilla 2 (fig., 10).—Outer plate the larger, covered with
exceedingly fine long hairs; expanded distally, bulging behind,
furnished with an apical cluster of stiff bristles, and a row of
sparsely plumose sete extending some distance down the inner
margin. The inner plate, distally narrowed, is provided with 2
rows of setz; one row, submarginal, consists of 20 long, delicate,
finely plumose set ; the second row contains an apical cluster of
stiff setae, with 8 inset along the margin, these latter of the same
structure as the plumose ones of the outer plate. In Grube’s
specimen the upper portion of the inner plate has been torn
away; the 2 apical setz represented in the figure 2 m‘ are two
of the plumose row of the outer plate.

Mawillipeds (figs. 11 & 12).—IJnner and outer plates well de-
veloped, but narrow. The apices of the inner plates are inset
with 4 long, feathered, setiform spines along the margin, with
a group at the inner angle of 4 long delicate plumose set
set together in a little hollow, 3 similar ones extending down
the inner margin, and 2 submarginally on the outer side. The
outer surface is longitudinally ridged. On this surface close to,
but just below, the inner angle, is a small cowpling-spine (fig. 12),
stout, and bent upwards. It appears to be tuberculated on its
inner side, but the detail is obscure, even with a ;4,th oil-immersion
lens. The ower plate carries a marginal row of flat feathered
spines (15 in Grube’s large specimen, 13 in Chevreux’s, 7 in the
smallest of all), the 2 apical ones much the longest, and sub-
marginally on the outer side a few long feathered set. The
2nd joint of the palp is the longest, in Grube’s specimen twice
the length of the Ist, furnished with numerous plumose sete ;
the 3rd is produced over the insertion of the finger as in the type
species; finger small, obtuse, carrying apically 1 long stout
dentate spine, 1 smaller one, and 2 sete in Chevreux’s specimen,
3 stout spines and 4 ‘sete in Grube’s, 1 spine and 2 sete in
Norman’s.

First Gnathopods (figs. 18, 15, 17, & 18).—2nd joint narrow at
the base, but widening distally; posterior margin convex, with
2-4 very long delicate setz inset midway; anterior margin
straight, carrying a row of delicate plumose setze, another row of
longer similar setz is found on the under surface as in the type
species ; 3rd joint bulging behind, fringed with numerous closely
set long feathered sete; 4th joint the smallest, also fringed
posteriorly with feathered sete shorter than those of the 3rd.
The 5th is considerably longer than the 6th, with 5-7 transverse
rows of plumose sete, and 5-7 clusters of rigid serrate bristles
along the hind margin. 6th oblong, about twice as long as broad,
with 4—5 transverse rows of sete onthe hind margin ; palm short,

582 MRS. E. W. SEXTON ON THE

truncate. The long spine on the hind margin in Chevreux’s
figure, (16) pl. xi. ig. 2a, is the longest of a group of 3 inset
on the under side of the hand, and projecting beyond the hind
margin. The palmar margin is mentioned by Chevreux (p. 91)
as affording a distinguishing character for the two forms—concave
for pectinatus, and convex for dellavallei,—but an examination of
a series of specimens shows conclusively that this difference is due
to sex. In the male (figs. 15 & 18) the margin is concave, and
the palmar angle forms a right angle with the hind margin;
in the female (fig. 17) the palmar angle is rounded, the
curve commencing at the insertion of the finger, and merging
imperceptibly into the hind margin. The palm is strongly
serrated in all; in the female the serrations turn the corner and
reach as far as the large spine Just referred to. The palmar spines
ave of the same structure as those of the type species. The finger
is more than twice the length of the palm (a little longer in pro-
portion in the young form), curved, with a strong auxiliary claw ;
a long, stiff, finely serrated spine and 1 setule are inset in the
notch, 1-2 setules behind the claw; the rest of the inner margin
is firmly serrated.

Second Gnathopod (figs. 19 & 20) as described by Grube,
Norman, and Della Valle. Figures of the finger, so characteristic
of the species, are given from Norman’s and Chevreux’s specimens
in order to show the identity of the two forms pectinatus and
dellavallet. Grube’s specimens agree exactly with these. In the
young form the finger is slightly longer in proportion to the pre-
ceding joint than in the older animals, not equalling quite half
its length in Norman’s specimens, and only about one-third the
length in Grube’s largest.

Pereopods | and 2 glandular, alike in structure ; 2nd joint large,
fringed posteriorly with long delicate sete, more numerous on the
2nd pereopod; anterior margins with from 4-6 similar sete:
Ath joint about half the length of the 2nd, slightly expanded
distally ; 5th joint narrower and shorter than 4th; 6th joint
slightly longer than the 4th; the two terminal joints tapering
gradually to the acute tip of the 7th; the 7th subequal to the 6th
in length, with the glandular aperture opening at the tip.

Hinder Pereopods (fig. 21) more strongly built than the pre-
ceding; rapidly increasing in length. Pereopod 3: 2nd joint
large, a little longer than broad, equalling in length the 3 fol-
lowing joints taken together; both margins convex, the anterior
beset at intervals with 7 small sensory spines, the first 3 un-
accompanied by sete, the 4th with 1 seta, the 5th with 2, 6th
with 3, and the distal one with 4; posterior margin produced
below in a rounded lobe, crenulate, 6 of the crenulations with
setules inset, the 2 crenulations on the lobe each with a ciliated
hair; 3rd joint short and broad, with a cluster of sete, 1 ciliated
hair, and 1 spine; 4th joint half the length of the 2nd, stout,
slightly expanded anteriorly, with the spines of the structure
peculiar to the hinder perzeopods and uropods (fig. 21); 5th joint

AMPHIPOD GENUS LEPTOCHEIRUS. 583

shorter than 4th, with two clusters of spines on either side;
6th about as long as the 4th, but much more slender, spinose,
with groups of spines on either side and | very long stout spine
inset behind the finger. Finger short, not half the length of the
preceding joint, much curved, with 2 specialised bristles towards
the apex, one on the inner side lying flat against the finger and
reaching to the apex, and the other on the outer side, setting out
at right angles to the finger. This one appears to have a flattened
tip, and the inner one to be feathered, but the detail is almost
impossible to see owing to their extreme tenuity and the angle
at which they are placed. Perzeopod 4 is noticeable for the great
length of the spines of the 5th and 6th joints; 2nd joint rounded
oval, produced behind into a rounded lobe, 8 spines along the
anterior margin, the distal 5 accompanied by increasing clusters
of ciliated hairs and sete; posterior margin with 10 crenulations,
and a submarginal row around the lobe of 5-8 ciliated hairs; 4th
and 5th joints practically subequal to each other in length, 5th
narrower than 4th; 6th considerably longer and more slender ;
the spines on this joint are longer than those on the 5th, 4 clusters
increasing in length and number, on either side, 2 of the clusters
behind the finger equalling the joint itself in length; they are
very brittle, several of the specimens not having a single perfect
one remaining. Finger lightly curved, of the same structure as
the finger of the preceding pereopod. Pereopod 5: 2nd joint
narrow proximally, widened distally ; the proximal end of the
posterior expansion is produced subacutely, the distal end, as in
pereopods 3 and 4, forms a rounded lobe; on the posterior
margin are 8 crenulations with a setule on each, and 20 ciliated
hairs in a submarginal row; the anterior margin has 8 spines as
in pereopod 4, and 3 long fine hairs proximally. The 4th—6th
joints rapidly increase in length and decrease in width ; 4th and
5th each beset with 3 groups of stout spines on either side;
6th twice the length of the 4th, with 5 groups of spines on
either side, the terminal posterior group of spines and setz of
extraordinary length. Finger as in pereeopod 4.

The smaller specimens agree in all details with this description,
the only difference being the lesser number of spines.

Pleopod 1 (fig. 22): peduncle short, hardly half the length of the
outer ramus, with about 10 long plumose setz on the outer side ; 2
small coupling-spines, and | long plumose seta inset together on the
inner. The outer ramus 12-jointed, considerably shorter than the
inner ; inner ramus 14-jointed, with 4 cleft spines. Pleopods 2
and 3 alike; peduncle more than half the length of the outer
ramus, with only 2 or 3 fine hairs. The rami are shorter than
in the lst pleopod ; outer ramus with 11 joints; inner with 13 ;
4 cleft spines in the 2nd pleopod ; 2 in the 3rd.

Uropods (figs. 23-26) extend backwards to the same level.
The peduncle of uropod 1 is shorter than the outer ramus, with
5 slender spines on the upper curve, and 3 inset diagonally midway
on the outer surface; the falciform apical process reaches to half

584 MRS. E. W. SEX'TON ON THE

the length of the outer ramus. The outer ramus is shorter than
the inner, both furnished with slender spines. In uropod 2 the
peduncle is almost as long as the outer ramus; outer ramus
the shorter; the spines are shorter and stouter than those of
uropod 1. I have figured uropod 3 in detail (figs. 23, 24, &
25), because it has been used by Norman (36, p. 89) as dis-
tinguishing the two forms. A comparison of the figures will
show the identity of structure. The outer ramus has a minute
apical joint carrying a long rigid spine, this joint being larger in
proportion in the young form; the detail of the spines in the
small specimen (fig. 23) could only be seen with the jth oil-
immersion.

The Telson (figs. 27 & 28) is of exactly the same structure in
all the specimens; lateral angles greatly elevated, each produced
to an acute point, each with 1 long spine and 1 short sensory
ciliated hair on the margin, and 2 ciliated hairs underneath. The
subacute apex curves upwards also, but not to the same degree
as the lateral angles. igure 27 is taken from Chevreux’s speci-
men ; in two of Grube’s, the tip was as represented in fig. 28, in
the third, the largest, it was as in fig. 27.

Colour.—Grube’s largest specimen still retains its vivid colour,
a yellowish tint, with markings in dark brown. The dis-
tribution of the pigment agrees perfectly with Costa’s figure (19)
pl. ui. fig. 8. The markings are composed of cloudy brownish
patches stippled thickly but irregularly with blackish-brown spots.
The dorsal part of the head is‘covered with dark brown pigment,
shading into black near the eyes, these dark patches causing
the eyes to appear larger than they really are. The lst perzon-
segment is plain; the 2nd has only faint, patches, one on the back,
and one on each side; the 3rd and 4th with their corresponding
sideplates, and the 5th are thickly covered with the pigment,
arranged in a band along the middle of the segment; the 6th is
plain ; the 7th and the first 3 pleon-segments with their epimera
are darkly pigmented ; the 4th pleon-segment has a faint patch
on each side; the rest of the animal is plain. The general effect
is very striking, the animal showing a dark head, 3 dark rings
anteriorly, and 4 farther back. The amount of colour probably
varies with the locality. | Walker, (48) p. 310, describes it as
‘“deep yellow, with transverse brown lines on all the segments.”

Distribution.—This species has a very wide distribution :—

SHETLAND Istus : Norman (84) p. 283, as Protomedeia pectinata ;
dredged; St. Magnus Bay, deep water.

Trish Sea: Walker (48) p. 310, as Leptocheirus pilosus ;
dredged; 17 fathoms.

W.&S. Coasts or IneLanp: Walker (50) p. 169, as Leptocheirus
pilosus; 40 fathoms ; bottom deposit, gravel.

CHANNEL Istanps: Walker & Hornell (49) p. 54; as Lepto-
cheirus pilosus Zadd.=L. pectinatus Norman; 7 fathoms;
bottom deposit, gravel, clinkers, and shells. Chevreux (16)

AMPHIPOD GENUS LEPTOCHEIRUS. 585

p. 91, as Leptocheirus pilosus ; tidal zone; bottom deposit,
rather coarse sand. Norman (37) p. 369, as Leptocheirus
pectinatus.

France: Ocranic Coasr: Chevreux (11) pp. 290, 309, as
Ptilocheirus pectinatus ; 10-19 m. ; bottom deposit, nullipores
and mud, Chevreux (15) p. 482, as Leptocheirus pilosus ;
10-20 m.

Bay or Biscay: ss. ‘ Huxley,’ 47° 48’ N.; 7° 46° W.; 109
fathoms.

MEDITERRANEAN : Chevreux (16) p. 91,as Leptocheirus fasciatus.
PROVENCE : coast of ALGERIA: Corsica, dredged in 12 fathoms:

Chevreux (17) p. 4, as Leptocheirus fasciatus.

Bay or Napues: Costa (19) p. 155, as Protomedeia fasciata.
Della Valle (21) p. 450, as Leptocheirus pilosus ; 10-20 m. ;
bottom deposit, sand.

Aprtatic : Grube (22) p. 73, as Protomedeia pilosa. Grube (24)
p. 403, as Protomedeia hirsutimana® and Pr. pilosa.

SENEGAL: Chevreux (in litt.).

Wasin, Brir. KE. Arrica: Walker (51) p. 341, as Leptocheirus
pilosus; 10 fathoms ; bottom deposit, mud.

LEPTOCHEIRUS BISPINOsUS Norman. (Plate XVIII. figs. 17-20.
Text-fig. 146.)

1866. Protomedeia hirsutimana Heller (25) pp. 34 & 35.

1893. Leptocheirus guttatus Della Valle (21) p. 430, pl. xii.
figs. 15-24.

1908. Leptocherrus bispinosus Norman (38) p. 308, pl. xii.
figs. 7-9; pl. xii. figs. 1-3.

The specimen described by Heller as Protomedeia hirsutimana
Sp. Bate is preserved in the Hofmuseum, Vienna. The de-
scription and figures of it, most kindly sent to me by Dr. Pesta,
prove it to belong to the same species as described by Norman
in 1908 under the name of ZL. bispinosus. -

The guttatus of Della Valle, in my opinion, must also be in-
cluded in this species. A comparison of the two accounts, Della
Valle’s and Norman’s, leaves no room for doubt. The only point
of difference is in the number of joints in the accessory flagellum :
Della Valle gives the number as 2, subequal in length, and as long,
taken together, as 2 joints of the primary flagellum; Norman
as 5, as long as 3 joints of the primary. It seems probable, aither
that Della Valle had a young specimen before him with only two
joints developed, or, what I think more likely, that the accessory
flagellum was broken. In the other specimens of this species (and,
indeed, in all the specimens of this genus that I have examined)
the terminal joint of the accessory flagellum is very small and
tipped with long sete, but Della. Valle says definitely that the 2nd
joint in his specimen was equal in length to the lst, and in his
figure he shows only 2 small sete instead of the usual long apical
cluster, I have added below some details to the description given

586 MRS. E. W. SEXTON ON THE

Text-fig. 146.

Leptocheirus bispinosus Norman.

The figures were drawn by Dr. Otto Pesta from Heller’s specimen
in the Hofmuseum, Vienna.

Fig. a. Superior antenna, accessory flagellum, left side, X 100.

Fic. 6. ee é 53 * right side, X 100.

Fig. ec. First pereopod, X 60. Fig. d. Hand of first enathopod, X 100.

Fig. e. Finger of second gnathopod, X 200. Fig. f Sideplates 2 and 3, x 60.

Fig. g. Pleon-segments 3-6, X 60.

AMPHIPOD GENUS LEPTOCHEIRUS. 587

by Canon Norman for the sake of a more complete comparison
with Della Valle’s account.

The species resembles guttatus Grube in many respects, the
principal distinguishing points
antennz ; the hand of gnathopod 1 ; sideplate 2 ; the shape of the
basal joints of the hinder peropods ; the armature of the pleon ;
and uropod 3

Description (taken from the type specimen of bispinosus, a
female, which Canon Norman kindly allowed me to examine) :—

Head \onger than pereon-segments 1 and 2 taken together ;
lateral corners rounded.

Hyes very large; pigment brownish black; ommatidia large.

Sideplate 1 free of sideplate 2, about half its width, but not
equalling it in length, a little expanded distally and fringed with
long setze. Sideplates 2-5 much as in the type species for shape.
The 2nd is the largest and deepest, much deeper than the corre-
sponding body-segment; expanded inferiorly, hind margin
straight, inferior margin rounded and fringed with numerous
plumose sete (cf. text-fig. 146, f, with Norman’s pl. xiii. fig. 2
and Della Valle’s pl. xii. fig. 17). The 3rd is deeper than the
4th ; and the 4th a little deeper than the 5th; all fringed inferiorly
with sete.

Pleon.—Segment 3, hind margin produced, rounded, crenulate ;
crenulations 8 in number, each with a setule; inferior margin
concave, furnished with numerous strong sensory spines sub-
marginally. Pleon-segment 4 as described by Norman, with ‘a
strong and acute angular backward projection on each side” ;
Dr. Pesta’s account agrees with this, “das 4. Pleonsegment hat 2
seitliche nach aufwiirts geschwungene ‘ angles’” (text-fig. 146, g) ;
Heller erroneously notes them as occurring on the 5th. The 5th,
indeed, carries 2 dorso-lateral angles (as stated by Della Valle),
but these are exceedingly small and difficult to see in situ, not
much produced, acute, each with a seta inset behind.

ANTENN (text- figure 146 a, b).—Superior Antenna, as figured
by Della Valle: 1st joint of peduncle shorter and broader than
the 2nd, carrying a stout spine on the inner distal angle; 3rd a
little more than one-third the length of the 2nd; primary
flagellum with 16 joints, each, from the 3rd, with a small sensory
filament ; Dr. Pesta says of Heller’ s specimen that the number of
joints on the right side is 17, on the left 18. The accessory
flagellum in Norman’s specimen extends to the 3rd joint of the
primary, 5-jointed, the first 4 joints subequal to each other in
length, the terminal one minute, tipped with long setze ; in Heller’s
specimen it does not reach to the distal margin of the 3rd joint of
the primary, ““4-gliedrig, jedoch ist das Endglied rechts kiirzere
als links.”

Inferior Antenna: Ath joint longer than 5th; flagellum about
the same length as the 5th, com posed of 7 joints, the first as long
as all the others taken together.

First Gnathopod (PI. XVIII. fig. 17): 3rd and 4th joints more
densely setose than in the other species, the plumose sete being

588 MRS. E. W. SEXTON ON THE

arranged in transverse rows in addition to the marginal fringe.
5th joint slightly longer than 6th; 6th twice as long as broad,
widening a little towards the palm. The palmar margin is convex,
its limit defined, as in the preceding species, by a long sensory
spine inset on the under surface; on the outer surface it carries
a submarginal row of about 10 strong sensory spines, and on the
inner side a thick row of setiform spines. The finger when closed
reaches to the large spine; it is curved, with a strong auxiliary
tooth subapically, and 6 smaller teeth on the inner margin, each
tooth with a setule beside it. In the notch near the apex 2 or 3
longer setules are inset.

Dr. Pesta’s description is as follows (text-fig. 146, d): “das
Handglied des 1. Gnathopoden ist vorne etwas verbreitert, die
Klaue so lang wie die konvexe Seite desselben ; der konvexe Rand
die ‘ Hand’ besitzt eine Reihe von kriiftigen Dornen, die gegen
die Klaue gerichtet sind.”

Second Gnathopod (fig. 18): 2nd jomt unusually long, equalling
in length the joints 3-6 taken together; 3rd, 4th, and 5th joints
subequal to each other measured along the inner side. The finger
is nearly straight, about half the length of the preceding joint ;
“leicht gebogen und spitz” (text-fig. 146, e).

Perceopods 1 and 2.—The length of the 4th joint forms a dis-
tinguishing character for this species. The peropods are about
the same length, but in perzopod | the basal joint is longer than
- in perzopod 2; and the 4th joint is longer in the latter than in the
former, being equal to the basal joint in length. Dr. Pesta says
“das 4 Glied des 1. Peraeopoden ist linger als das 6 Glied und fast
so lang wie das 2 Glied.” (Cf. text-fig. 146, ¢, with Norman’s pl. xii.
fig. 8 and Della Valle’s pl. xii. fig. 24.) The 2nd joint carries
posteriorly numerous delicate sensory set, some of great length ;
4th joint long, of equal breadth throughout; 5th about half as
long as the 4th; 6th longer than 5th, both with clusters of long
sparsely feathered sete posteriorly ; finger shorter than the 6th,
subequal to the 5th in length, with the glandular aperture
opening at the tip.

Hinder Perceopods (fig. 19) as figured by Della Valle. In perzeo-
pod 3 the basal joint has the posterior margin straight and
crenulate ; in pereopod 5 the posterior expansion of this joint is
much wider distally than at the base, and the margin is rounded
and serrate, the lower portion carrying numerous plumose sete.
Fingers curved and falciform. In Heller’s specimen, perzeopod 3
is missing, but Dr. Pesta says of the last two pairs “die End-
klauen sind einfach (nicht ‘ bifid’),” as in Spence Bate’s hirsuti-
mana. In Norman's figure, pl. xii. fig. 9, the terminal joints
are lying in such a position as to quite conceal thespines. I have
therefore refigured them to show the armature.

Uropods 1 and 2 (text-fig. 146, g) much as in gutiatus Grube ;
very spinose, the spines of uropod 2 shorter and much stouter than
those of uropod 1. The falciform process of the peduncle reaches,
in uropod 1, to the tip of the outer ramus; in uropod 2 beyond
the tip. The inner rami differ from the other species in having

AMPHIPOD GENUS LEPTOCHEIRUS. 589

slender, feathered set, or setiform spines, in addition to the other
spines. These are found on the under margin, 3 in uropod 1,
short, inset at intervals, and 2 clusters of long ones in uropod 2,
near the apex. In uropod 3 (fig. 20) the outer ramus is fur-
nished with an apical cluster of 5 spinesand 3 or 4 long feathered
setee similar to those of the 2nd uropod, and 1 cluster of small
spines midway, in Norman’s specimen (2 clusters figured by Della
Valle, probably from an older specimen) ; the inner ramus is tipped
with 1 stout spine and | seta, with 1 spine midway, in Norman’s
specimen (2 in Della Valle’s).

Telson.—Thé apical margin of the telson is represented in Della
Valle’s figure as truncate, but, as in guitatus Grube, this appear-
ance is due to the angle at which it is carried. The mar gin 1s
really convex, but as in the natural position it is a little bent i in
under, the convexity is not apparent from the dorsal view.

Colour given by Della Valle as greyish yellow without markings ;
by Dr. Pesta as “ gelbbraun, Auge schwarz” ; but, he adds, “ diese
Farbung diirfte nicht natiirlich sein.”

Distribution :—

Bay or Biscay: Norman (38) p. 309, as ZL. bispinosus,
depth 35-60 fathoms.

Narues: Della Valle (21) p. 482, as L. guitatus ; bottom deposit,
corallines.

Apriatic at Lesina: Heller (25) p. 35, as Protomedeia
hirsutimana.

Gutr or Bone and Coast of Tunis: Chevreux (18) p. 3, as
L. bispinosus.

The only details given by Catta concerning his species massili-
ensis, viz. the nature of the ground on which it was captured
“fonds coralligénes,” and the colour, “ un beau jaune, tandis-
que l’ceil était completement noir,” would seem to point to
L. bispinosus.

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« 42%

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(44) Sressine, T. R. R.—Das Tierreich. 21. Lief. Amphipoda.
I. Gammaridea. Berlin, 1906, pp. 625-630.

(45) Srimpson, W.—Synopsis of the Marine Invertebrata of Grand
Manan. Smithsonian Contributions to Knowledge,
1853, vol. vi. no. 5, pp. 55-6.

(46) VERRILL, Smrru, & Harcer.—Catalogue of the Marine In-
vertebrate Animals of the Southern Coast of New England
and adjacent waters, by A. E. Verrill, 8. I. Smith, and Oscar
Harger. U.S. Commission of Fish and Fisheries, 1873,

. 561.

(47) Ween A. O.—The Amphipoda of Bate and Westwood’s
‘ British Sessile-eyed Crustacea.” Ann. & Mag. Nat. Hist.
(6) vol. xv. 1895, pp. 469-470.

(48) WALKER, A. O.—Revision of the Amphipoda of the L.M.B.C.
District. Trans. Liverpool Biological Society, vol. ix. 1895,
pp. 310, 311.

(49) WaLker & Horneti.—Report on the Schizopoda, Cumacea,
Tsopoda, and Amphipoda of the Channel Islands. Journ.
Marine Zoology and Microscopy, vol. ii. no. 7, 1896,

. 54,

(50) ae A. O.—Malacostraca from the West Coast of
Treland. Trans. Liverp. Biol. Soc. vol. xu. 1898, p. 169.

(50a) WALKER, A. O.—Malacostracan Fauna of the Mediter-
ranean. Journ. Linn. Soc. vol. xxviii. no. 12, 1901,
p. 305.

(51) Waker, A. O.—Amphipoda Gammaridea from the Indian
Ocean, Brit. East Africa, and the Red Sea. Trans. Linn,

Sce. vol. xii. pt. 4, 2nd series, Zool. 1909, p. 341.

AMPHIPOD GENUS LEPTOCHEIRUS. 593

(52) Waker, A. O.—Notes on Amphipoda. Ann. & Mag. Nat.
Hist. (8) vol. vi. 1910, p. 33.

(53) Zappacu, E. G.—Synopseos Crustacecrum Prussicorum
prodromus. Kéonigsberg, 1844, pp. 8, 9.

(54) Zappacu, E. G.—Die Meeres-Fauna an der preussischen
Kuste. Erste Abth. Schrift. d. Phys.-dkon. Ges. z. Konigs-
berg, 19. Jahrg. 1878, pp. 18, 19.

EXPLANATION OF THE PLATES.

PuatE XVII.

Leptocheirus pilosus Zaddach.

Fig. 1. Whole animal, Zaddach’s specimen, young SLR eats SEO aE Ne EAD
2. Superior antenna, seen trom above, young 2. Zaddach’s specimen. X ASO.
8. Part of inferior antenna............... eves i 3 X 50.
AM Wippersliplscrnsaesctexcgeee meee. basal ss ¥3 53 x 95.
Belhowertipissn5-eu eeu emer rcre. mane ces ue * XK 98.
Gap Man dibleped saat res eae Wee ae ene ee le 5 s x 95.
Teale xall aoe carte cara aie cence Mean Fess PH “i x 95.
Sat Texall Ate cea eet seacecasta coe cate ete i “ x 95.
Oro Mila xallamed sisunchescasedceteomemadteeeveri las ta. 95 Pa x 95.

10. First gnathopod .. SASS os 35 Xx 50.
11. Hand of the tirst enathopod. 9 . Norman’s POOVNIN coconsassconcen >< (0)
12. Gin (Chevréuxisy 8 os.) 4 ean ae x 50.
13. Finger of first enathopod. Large 2 3 5 x 95.
14, X 95.

3 ; 5 apni secaeces
15. Serrate bristle from 5th joint, first gnathopod. Large 3.
Chevreux’s spec x 175.

16. Second gnathopod. Young ie . Zaddach’s specimen ..... eae Oxo!
17. Finger of second gnathopod. Young 2. Zaddach’s specimen anes DLO!
18. 5 = PA 9. Norman’s Pen aces x 140.
19. First pereopod. Young 2. Zaddach’s AYNSOUNEN “UGescusosootace: éoacco, 4 GAO):
20. Second - ieee a "3 Rr om ieereatinae or <5) 0)5
21. Third Bn saedess 33 Say i uy, ates atrckoperynatna carseat x 50.
22. Fitth ae 55 55 adsisasdaacka eee 2 DADO!
23. Fifth pereeopod, under surtace. Large g. Chevreux’s specimen... X 40.
24. Upper coupling-spine, first pleopod. Young 9. Zaddach’sspecimen. X 290.
25. Telson and third uropods, dorsal view. ,,_,, 53 x 60.
26. Telson, lateral view. Young 9. Zaddach’s specimen . ise kVOO!

Poate XVIII.

Fig.1. Whole animal. Grube’s type specimen ... Leptocheirus guttatus. X 20.

2. Accessor y flagellum, right antenna. Nor-
man’s specimen .... atieeh. on 2 x 76.

3. Accessory flagellum, left antenna. Same
specimen ..... > x 75.

4, Accessory flagellum, right antenna. Another
SPECIMEN On erence neon crtie miea Nene = ss x 78.

5. Hand and finger, “first sgnathopod. Chev-
reux’s specimen, x 7bs

6. Finger of second enathopod. “ Chevrenx’s
specimen,? ... 3 3 x<a7ios

7. One of the smaller apical spines, uropod 2.
Norman’s specimen .... 55 5 X 265.

8. Third uropod, drawn in “situ. Grube’s
speciinen ... j PS x 76.

9. Apical margin of telson, ; a little upturned.

Grubesispecimentimyper ese meena cern 5 ¥ x 75.

594 ON THE AMPHIPOD GENUS LEPLOCHEIRUS.

Fig.10. Hand of first gnathopod, ¢, looking down

on the hind margin, from a ae from

Vineyard Sound . : sab . Leptocheirus pinguis. X 22.
11. Finger of second onatbopod, s same - specimen. a5 4 * 27.
12. Second joint, without the bristles, uropod 3,

QD, Wimesyemrdl SOWIE soo scocaseseosnosbe sn caneae ; % x 75.

13}, Finger of first gnathopod, small specimen,
4 mm., from Banff ........ : ....... Leptocheirus hirsutimanus X 58.
14. Tip of finger, second enathopod, small speci-
men, 4 1mm., from Banft.. Wenn:
15. Finger of third per: eopod, arge specimen,
10 mm., Shetland Isles, 1867 .................. 3 ep x 146.
16. Outer ramus, third uropod, small specimen,
Avmntn.; IB anittik tee eee Leen eee
17. Hand and finger, first pais aaa oF
Norman's type : specimen : Me
18. Finger of second enathopod, @ ‘Norman’s
type specimen ...... x
19. Terminal joints, fifth “peraopod, o, ~Nor-
man’s type specimen ........ : n 55 x 28.
20. Uropod 3, 9. Norman’s ty pe ‘specimen — x

» » x Mo.

a op < 58.

. Leptocheirus bispinosus. X 50,

be 99

32 »

PratEt XIX.

Leptocheirus pectinatus Norman.

Fig 1. Accessory flagellum, superior antenna. Norman’s specimen, 6 ...... x 1465.
2. i 3 33 Fs Chevreux’s specimen, large @. X 75.
3. ut * Py is Grube’s largest specimen, 2. X 75.
fe Wipper lip) eeeeaee tenes oe eeeen ee eeee a Oh evreuxes) specimens lansiend) sn Guor
SH ILOW EL: Niple cereale: ees eee ee Renee 5 op Se, Oe:
6. Mandible 35 3 hehe ate 1S 78.
7. Manilla 1 : 3 ‘ re eS, ae
. Biturcate spine, outer plate, maxilla 1. es aS as son
9. Dentate spine, 58 ” » 2 ) oy) ” x 435.

10), Wlesalllle Fo. cs0cs0 0: a 3 RT SES
11. Mawillipeds ........ HA be SAT Paso OM 00s
12. Coupling-spine, mner : plate, maxillipeds. 5 » x 430.
13. First gnathopod (the feathering of the setze on joints Be 6 not ‘show n).
Norman’s specimen, young ¢,2°5mm. X 9d.
14. First sideplate............. “5 young 9,275 mm. X 95.
15. First enathopod. “Chevreux’s specimen, large 6, 5:25 mm............. x 40.
16 Hirst sideplate. Giube’s specimen, large 9) ...)... x 40.
17. Finger and palm, first gnathopod. Grube’s specimen, large 9 ...... x 95.

(The other sete omitted in order to show the palmar spmes clearly. )

18. Finger and palm, first gnathopod. Chevreux’'s specimen, large g... X 95.
19. Finger of second gnathopod. Norman’s specimen, 2 .................. X 290.
20. ‘ 4 Chevreux’s specimen, 6... Agencies. WOM:
21. Sensory spine, 6th jomt, perseopod 4. ,, es atencdactocassaidanae | A ek)
22. Tip of 4th cleft spine, pleopod 1. as a Rt ied” oh Sle aa al X 300.
23. Third uropod. Norman’s specimen, young 2, 2° 75 mm. MY, BRO EE OC ZO0F
24. iy ap Chevreux’s specimen, large Sq DED WAV, ooo soncoooscnes x 95.
25. _ Grube’s specimen, large Q acae botenehmans | Oe
26. U ropods and telson. Chevreux’s specimen, large Be “6mm... x 40.
27. Apex of telson. an 3 A 35 SR x 58.

28. 55 5 Grulbeisispecimen. Oe volo amms ss eeee eee Doe

ee eet

PAS, Leah IPL, 2OX,

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y

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West, Newman lith.

OSTRACODA FROM MADEIRA.

PAS. isi, Ill, SOC.

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OSTRACODA EPROM MADEIRA:

fetidion.

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ee

ah

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OSTRACODA FROM MADEIRA.

ON MARINE OSTRACODA FROM MADEIRA. 595:

27. Notes on Marine Ostracoda from Madeira. By G.
Srmwarpson Brapy, M.D., LL.D: D.Se., F.B.S.,
C.M.Z.S.

fReceived January 13, 1911: Read March 21, 1911.!
(Plates XX.-XXIL*)

T am indebted for the specimens which form the subject of the
following notes to my friend the Rev. Canon Norman, F.R.S.,
by whom they were collected in the spring of the year 1897.
The mountings submitted to me consisted altogether of dried
shells, and in no case was the contained animal preserved,
excepting in a few of those collected between tide-marks,
and only in some of these littoral forms was the original colouring
discernible. Apart from such as appear to be new to science,
the collection is interesting as extending the known range of
several species from the European and North Atlantic areas much
further southward, though not quite into the tropical zone. — All
the species belong, so far as can be ascertained from the shell-
characters, to well-known genera. The following lists show
separately the species found in the littoral zone and in deep
water.

Between tide-marks.

}

Cythere convexa Baird. Cytherura maculosa, sp. n.

eS albo-maculata Baird. 55 cellulosa Norman.
Xestoleberis depressa G. O. Sars. | Sclerochilus levis? G. W. Miller.
5 nigromaculata, sp. n. (Pl. XXII. fig. 10.)
ss margaritea Brady. | Paradoxostoma hibernicum Brady.
Loxoconcha impressa Baird. 59 arcuatum Brady.
Dredged in 30-70 fathoms.
Bythocypris reniformis Brady. | Loxoconcha obesa, sp. n.
Macrocypris decora Brady. | D decipiens G. W. Miller.
Pontocypris succinea G. W. Miller. sf impressa Baird.
35 dispar? G. W. Muller. | be? subalata, sp. 0.
Bairdia amygdaloides Brady. Xestoleberis latissima, sp. n.
» obtusata G. O. Sars. | Cytherura striata G. O. Sars.
» dubia, sp. n. | Bs cribrosa, sp. n.
> mediterranea G. W. Miller. | 5 eribriformis G. W. Miller.
» acanthigera Brady. | a fossulata, sp. n. |
Argillcecia atfinis, sp. n. Cytherideis subulata, var. crenulata
Cythere crispata Brady. Brady.
» cingulata, sp. n. Sclerochilus contortus Norman.
» tuberculata G. O. Sars. As levis? G. W. Miiller.
» emaciata Brady. (Pl. XXII. fig. 10.)
Cythereis runcinata Baird. Paradoxostoma gracile, sp. n.
» deformis, sp. n. fi flexuosum Brady.
» antiquata Baird. 4 cylindricum G. W.
>» jones Baird. Miller.
Cytheridea elongata Brady. Sarsiella capsula Norman.
Eucythere prava Brady & Robertson. Cytherella ? ovalis, sp. n.

* For explanation of the Plates see p. 601.

396 DR. G. S. BRADY ON

BairDIA DUBIA, sp. n. (Plate XX. figs. 1, 2.)

The outline of the shell as seen laterally is rhomboidal (fig. 1),
the greatest height situated near the middle and equal to half
the length; anterior extremity obliquely subtruncate, posterior
produced below the middle into a wide, obtuse beak; dorsal
margin forming a flattened arch, inferior nearly straight, gently
sinuated in the middle: seen from above (fig. 2) oblong, ovate,
twice as long as broad, greatest width in the middle, sides gently
curved, subparailel, extremities produced, the anterior obtusely
rounded, posterior prominent and submucronate. Surface of the
shell smooth, covered throughout with very small, closely-set,
impressed, circular punctations. Colour yellowish grey. Length
0°65 mm.

One specimen only, dredged in 30 fathoms.

ARGILL@CIA AFFINIS, sp.n. (Plate XX. figs. 9, 10.)

Shell, seen laterally, elongated, siliquose (fig. 9), greatest height
in the middle, equal to more than one-third of the length ;
anterior extremity evenly rounded, narrow, posterior scarcely
at all angulated ventrally; dorsal margin forming a continuous
even arch “throughout, ventral nearly straight through its whole
length: seen from above (fig. 10) the outline is narrowly subovate,
thrice as long as broad, greatest width in the middle, tapering
towards the subacuminate extremities, but more abruptly behind
than in front. Surface of the shell perfectly smooth. Colour
light grey. Length 0°54 mm.

Several specimens dredged in 70 fathoms.

The differences in form of shell between the various species of
Argillecia hitherto described appear to be extremely slight, but
that here figured does not seem fairly referable to any one of
them.

CYTHERE CRISPATA Brady. (Plate XX. figs. 3, 4.)

Specimens of C. crispata, an extreme form of which is here
figured, were dredged in 70 fathoms. This form differs from the
type in having the extremities—and to some extent the ventral
margins—irregsularly dentated. This condition, however, is found,
though to a much less marked degree, in some of the type-
specimens.

These specimens seem indistinguishable from a species described
by G. W. Miiller—C. diffusa*; perhaps also from C. elegans of

the same author.

CYTHERE CINGULATA, sp.n. (Plate XX. figs. 5, 6.)

Shell, seen laterally, oblong, subquadrate (fig. 5), of nearly equal
height throughout, height equal to at least half the length ;
extremities well rounded, the posterior rather the narrower, dorsal
margin nearly straicht, inclined gently from before backward,

* Die Ostracoden des Golfes von Neapel, p. 354.

MARINE OSTRACODA FROM MADEIRA. 597

ventral slightly sinuated in the middle: seen from above (fig. 6)
the outline is oblong, with straight, parallel, lateral margins and
strongly produced extremities ; the margins converge steeply in
front terminating in a stout median process, behind they converge
almost rectangularly and form a very wide central hump. Sur-
face of the shell covered with rounded and closely-set impressed
pittings, and bordered in front and behind with a smooth
depressed fillet. Length 0:78 mm.

One specimen dredged in 30 fathoms, and one in. 70 fathoms.

CYTHEREIS DEFORMIS, sp.n. (Plate XX. figs. 7, 8.)

Shell, seen laterally, oblong, subquadrate, greatest height
situated near the front and equal to nearly halt the length (fig. 7) ;
anterior extremity well rounded and bordered with a series of
short, blunt teeth ; posterior extremity much narrower, truncated,
prominently angular in the middle and divided below the middle
into a few irregularly tooth-like processes; dorsal margin sloping
gradually from the front backwards, ventral margin almost
straight: seen from above (fig. 8) the outline is compressed,
oblong, about thrice as long as broad, the lateral margins very
irregular, with a sharply prominent angle at the posterior third ;
extremities much produced, broadly truncate, with irregularly
dentate margins. Shell- surface irregularly rugose, with a wide
depressed band stretching round and within the anterior margin ;
a sharply elevated crest running obliquely across the posterior
half and terminating in a sharply produced angle on its dorsal
aspect. Length 0°5 mm.

Dredged in 70 fathoms.

It is impossible, by the shell-characters alone, to separate clearly
the species belonging to the two genera Cythere and Cythereis as
they are now under Stood by most authors, nor, as it appears to
me, are the distimetive characters of the contained animal much
more satisfactory.

LoxoconeHa OBESA, sp.n. (Plate XXI. figs. 3, 4.)

Shell, seen laterally, elliptical, greatest height situated m the

middle and equal to half the length (fig. 3); extremities evenly
rounded, the Cala ior somewhat the narrower of the two; dorsal
margin very slightly arcuate, ventral almost straight, not at all
sinuated : seen from above (fig. 4) the outline is very br oadly oval
with strongly produced mucronate extremities, width considerably
exceeding half the length, lateral margins very strongly convex.
Shell-surface perfectly smooth. Colour grey. Length 0°46 mm.

Dredged in 70 fathoms ; one specimen only.

LoxoconcHA DECIPIENS G. W. Miiller. (Plate X XI. figs. 1, 2.)

Several specimens, agreeing very accurately with the figures

given by Dr. G. W. Miiller, were dredged in 70 fathoms. One

quite characteristic specimen is figured here.

598 DR. G. 8. BRADY ON

LoxoconcHA SUBALATA, sp.n. (Plate X XI. figs. 5, 6.)

Shell, seen from the side, oblong, subrhomboidal, twice as long
as broad (fig. 5); anterior extremity obliquely rounded, posterior
very obliquely rounded below the middle ; ventral margin rather
deeply sinuated in the middle, curving suddenly upwards behind,
dorsal margin perfectly straight : seen from above (fig. 6) hastate
in outline, prominently angulated behind the middle, from which
point the sides converge with a gentle curve to the anterior
extremity, which is sharply acuminate; behind the two lateral
angles the margins converge rather sharply in an irregularly
sinuous curve to the posterior extremity : the general contour is
thus made of two wedge-shaped portions—an anterior larger
wedge and a posterior small one. Shell-surface rough, marked
by closely-set small fossze, sharply elevated in the postero-ventral
regions, beneath which it is depressed, forming a somewhat
flattened curved lip. Length 0°38 mm.

Dredged in 70 fathoms.

XESTOLEBERIS LATISSIMA, sp.n. (Plate XX1. figs. 10-13.)

Shell, seen laterally, oblong, subovate, quite twice as long as
broad (fig. 10), highest behind the middle; anterior extremity
narrowly rounded, posterior sloping with a steep curve to the
ventral margin, where it forms a rounded angle; dorsal margin
forming a somewhat flattened arch, sloping steeply behind, more
gently in front, ventral margin nearly straight: seen from above
(fig. 11) the outline is excessively tumid, ovate, widest behind the
middle, width equal to two-thirds of the length, mucronate in front,
broadly rounded behind; the end view (fig. 12) is very broadly
wedge-shaped, widest ventrally where the angles are moderately
rounded off, height equal to about two-thirds of the width. Shell-
surface perfectly smooth, marked in some cases with a very few
small papilliform tubercles. Colour white. Length 0°75 mm.

Dredged in 70 fathoms.

The outline shown in fig. 13 may perhaps be referable to the
male, the more tumid outline to the female.

XESTOLEBERIS NIGROMACULATA, Sp. n. (Plate XXII. figs. 1-3.)

Shell of the female, seen laterally, oblong, subreniform, greatest
height situated in the middle and equal to half the length (fig. 1) ;
anterior extremity depressed, rounded, posterior much wider and
evenly rounded; dorsal margin boldly arched, ventral sinuated in
the middle: seen from above (fig. 2) ovate, gradually tapering to
the anterior extremity, which is rather sharply pointed, much
broader and well rounded behind, width and height equal. Sur-
face of the shell smooth, yellowish in colour, with irregular
clouded dark patches, and bearing a few very minute distinctly
scattered circular papiile. Eye-spots very conspicuous. Length
0:55 mm.

Shell of the male (fig. 3) rather smaller, more markedly
depressed in front and more elevated dorsally.

Several specimens taken between tide-marks.

Caen tne

MARINE OSTRAGODA FROM MADEIRA. 599

Cy?HERURA MACULOSA, sp.n. (Plate XXII. figs. 6, 7.)

Shell, seen laterally, oblong, subreniform, highest in the middle
(fig. 6), height equal to half the length ; anterior extremity evenly
rounded, posterior produced in the middle into a wide obtusely
pointed beak; dorsal margin evenly arched throughout, ventral
rather deeply sinuated in the middle, prominent behind, thence
sloping with a sinuous curve up to the beak: seen from above
(fig. 7) oblong, three times as long as broad, lateral margins
parallel throughout the greater part of their length, converging
towards the front; anterior extremity broad and truncated,
posterior produced into a wide median beak. Surface of the
valves marked with faint longitudinal striz and with minute,
closely-set, impressed pits ; in most cases the middle of the valve
bears a dark transverse blotch of irregular shape, but this may be
absent or much reduced in size. Length 0°39 mm.

Several specimens from low-water mark.

CYTHERURA CRIBROSA, sp. n. (Plate XXII. figs. 4, 5.)

g, rhomboidal, height searcely equal
to half the length ; ainkierslert extremity well rounded, posterior
produced above the middle into a very large and rather sharp
beak (fig. 4); dorsal margin straight throughout almost its whole
length, ventral rather deeply sinuated in the middle, behind
which it bulges ventrally: seen from above, oblong, subovate,
width scarcely equalling half the length (fig. 5), rather abruptly
tapered and subacuminate in front, posteriorly abruptly truncated
and produced into a large central beak. Surface of the shell
honeycombed with large subrotund fosse, which are arranged in
irregular longitudinal rows. Length 0- 44 mm.

Many specimens dredged in 50 fathoms.

Shell, seen from side, oblong

CYTHERURA FOSSULATA, sp.n. (Plate XXII. figs. 8, 9.)

Shell, seen laterally, oblong, rhomboidal, of nearly equal height
throughout, length equal to more than twice the height (fig. 8) ;
anterior extremity obliquely subtruncate, often indented below
the middle, the sinuations bounded by two small nodules, two
rounded tubercles near the anterior extremity, just within the
superior margin, over the region of the eyes ; posterior extremity
oblique, produced above the middle into a large upward-pointing
beak; dorsal margin perfectly straight, ventral nearly straight,
with a slight ventral convexity: seen from below (fig. 9) the
outline is that of an arrowhead, the lateral acute prominences
situated near the posterior third, the width at that point equal to
more than half the length of the shell, behind these angles the
shell is suddenly narrowed, forming a rounded hinder end which
terminates ina lar ee sme en beak : " anteriorly the lateral margins
converge with a deep curvature to the front, ending in a wide
bluntly rounded extremity. The surface of the shell is variously
and very irregularly marked with furrowed undulations more or
less transverse in their direction, and by a conspicuous curved
longitudinal evest which ends behind the middle near the ventral

600 ON MARINE OSTRACODA FROM MADEIRA.

margin in a sharply angular projection: in some cases there is
a distinct longitudinal striation, more especially on the ventral
surface, and posteriorly near the base of the beak may be seen a
series of four nodules, the terminations of small curved carine
(fig. 9). Length 0-46 mm.

Several specimens were dredged in depths of 50-70 fathoms.

The surface-ornament of this species varies very much: the
foregoing description should be taken as belonging to a rather
strongly marked specimen. The variations of sculpture seem to
depend chiefly on conditions of age and sex.

CYTHERIDEIS SUBULATA, Var. CRENULATA, nom. n. (Plate X.XI.
fig. 7.) ,

Specimens, of which a figure is here given, seem to be identical
with a form deseribed in 1874 by myself and the late Dr. Robert-
son under the name Cytherideis subulata var. fasciata, the
varietal designation having been used on account of the presence
of a dark band across the shell. But the type specimens, taken
among the Scilly Islands and now in my collection, do not now
exhibit any such marking. It is possible that the markings may
have disappeared with exposure to the air, and as the published
name is inappropriate, I propose to substitute the varietal term
crenulata. In the Madeira specimens, which were taken in a
depth of 70 fathoms, the anterior crenulations are much more
developed than in those from Scilly, but in all other respects those
from the two localities are alike.

PARADOXOSTOMA GRACILE, sp. n. (Plate X XI. figs. 8, 9.)

Shell, seen laterally, oblong, subclavate, height equal to more
than one-third of the length ; extremities rounded off, the anterior
narrower and somewhat depressed (fig. 8), dorsal margin evenly
but not very strongly arcuate, ventral almost straight : seen from
above (fig. 9) extremely compressed, widest in the middle and
tapering evenly to the extremities, which are very acutely pointed,
width equal to about one-fourth of the length. Shell-surface
perfectly smooth and colourless. Length 0°55 mm.

One specimen only, taken in 70 fathoms.

CYTRERELLA (2) OVALIS, sp. n. (Plate XXII. figs. 11, 12.)

Shell, seen laterally, subelliptical. about twice as long as broad
(fig. 11); anterior extremity rounded above and below, almost
subtruncate, posterior narrower and somewhat oblique, dorsal
margin almost straight, ventral straight, upcurved toward the
posterior extremity : seen dorsally the outline is ovate (fig. 12),
twice as long as broad, greatest width behind the middle;
extremities well rounded, the anterior much narrower than the
posterior. Shell-surface perfectly smooth. Colour white. Length
0-42 mm.

Dredged in 70 fathoms.

PS SiS PE esi’

West, Newman chr.

3 CEE @ Uae

el eye Wide Tea

(Aisa ON ay

toy apevallte sulle iy) RV

West, Newman chr.

1. DARK GRIZZLE. Aan NATALIE TSH (Ey ICI, Mala
34 (ETRE A Ae aD) \Oella@NOMa IR.

PZ.S.1944. Pl. XXV._

wiles

West, Newman chr.

dy SDWVNIBNCe VATS. oy RINSE) WN TENE
oye) WORDT IMIR

PAS. 191 Pl Oa

West, Newman chr.

FEATHERS SHOWING PATTERN MARKINGS.

ON COLOUR INHERITANCE IN PIGEONS. 601

EXPLANATION OF THE PLATES.

Prate XX.

1, 2. Bairdia dubia. X 84.

3, 4. Cythere crispata. 110.
SO. 5 cingulata. X 110.
7, 8. Cythereis deformis. X 84.
9,10. Argillecia affinis. X 84.

EFratr XXI.

1, 2. Loxoconcha decipiens. X 90.
3, 4. 5) ohesa. X 84.
5, aa subalata. X 115.
7. Cytherideis subulata, var. crenulata. X 84.
Figs. 8, 9. Paradoxostoma gracile. X 84.
3. Xestoleberis latissima. X 110.

Puatre XXIT.

Figs. 1-3. Xestoleberis nigromaculata,? g. X 84.
4, 5. Cytherura cribrosa. X 100.
@, We 3 maculosa. X 100.
Saas - fossulata. .X 100.

Fig. 10. Sclerochilus levis? X 84.

Figs. 11, 12. Cytherella ovalis. X 100.

28. On Colour and Colour-pattern Inheritance in Pigeons.
By J. Lewis Bonsore, M.A., F.L.S., F.Z.8., and F. W.
Smauuey, F.Z.S.

[Received February 4, 1911: Read March 21, 1911.]
(Plates XXII-XXVL-*)

The following is a preliminary account of some experiments
undertaken by the authors to throw some light on the inheritance
of colour and colour-pattern in Pigeons. These experiments are
being continued, as the matter is a long and complicated one and
will of necessity take several years to complete.

The experiments on certain colours and colour-patterns, how-
ever, have been practically completed, and the results are briefly
given below.

Apart from the practical knowledge given in books on fancy
pigeons, no serious work has been published on the inheritance
of colour in Pigeons except Darwin’s (Animals and Plants under
Domestication, vol. 1. p. 197 et seg., 1868 ed.), and a recent paper
by Mr. Staples- Browne (P. Z. 8. 1908, p. 67).

The information in the ‘fancy’ books, valuable as a guidance for
practical breeding, is of little utility from the scientific point
of view, as details of pedigrees are often lacking. The work of
Darwin we have also had to pass over, for the present, owing to
the difficulty of making out with any accuracy the exact colour of
his birds from the terms he used. It is by no means intended to
imply imaccuracy in that most accurate of observers, but the
‘faney’ terms for colours, though well understood by breeders, do

* For explanation of the Plates see p. 619.

602 MESSRS. J. LEWIS BONHLOTE AND F. W. SMALLEY ON

not readily admit of a scientific interpretation unless the birds
themselves can be seen; frequently the same colour (from a scier-
tific point of view) may be known by different names according to
the particular breed of pigeons that may be under discussion.

Darwin’s experiments related chiefly to “reversion,” 7. e. the re-
appearance of the blue colour when distantly related breeds of
pigeons were crossed, and although our work has not been on the
same lines, the study of the blue colour should, none the less, bear
out the results arrived at years ago. We are not unmindful of
this, and for that very purpose hepe to mate up several pairs to
test Darwin’s conclusions.

Lastly there is Mr. Staples-Browne’s recent paper, in which he
has attempted to repeat Darwin’s experiments. This paper we
have been through very carefully, and find that in the main it
agrees with our results, but one or two little difficulties have to be
met, such as the occurrence of a Dun in Exp. 27, the absence of
Blacks in Exps. 9 & 11, the large proportions of white in
Exps. 16-23 and 30, and the occurrence of white feathers on a
homozygous blue.

We have no doubt that further work will clear up these slight
difficulties, which do not, however, greatly affect the main result ;
in fact a possible explanation of some of them has already suggested
itself to us, but the discussion of these is best deferred until the
results of our matings on the same lines as Darwin and Staples-
Browne have been obtained.

The majority of the birds used have been highly bred Dragoons,
but in a very few cases a Homer cross has been introduced.

The characters dealt with in this paper are :—

(i) Colour-patterns, ¢, e. Chequering, Grizzle, and Mealy.

(ii) Colours, i.e. Blue and Silver, with White and Red in those

eases where it is connected with Grizzle and Mealy.

Before detailing the experiments, however, it is necessary to
have a clear understanding of the terms used.

(i) Chequering. This is a pattern chiefly confined to the wing-
coverts, in which each chequered feather has a hght coloured
V-shaped patch at its distal end, the apex of the V being
nearest the base of the feather (see Pl. XXVLI. fig. 1).
The general appearance of a good chequered bird is shown
in Pl. XXIII. fig. 3, but it must be understood that
scientifically and in the experiments we have carried out,
a bird has been considered as chequered when it showed
the characteristic markings on its wings.

(ii) Grizzling. A grizzled feather is one in which the barbs are
partially white and partially coloured. This pattern is not
restricted to any particular part of a bird, but grizzled
feathers may be found in any feather tract including the
remiges and rectrices.

On a grizzled bird (.e.a bird with grizzled feathers)
whole coloured feathers are generally found as well as
feathers splashed with white (Pl. XXIV. figs. 1 & 2;

=a,

COLOUR INHERITANCE IN PIGEONS. 603

Pl. XXVI. figs. 2 & 4). The term ‘Grizzle’ is restricted
in the ‘fancy’ to birds whose pigment is Blue grizzled with
White.

(iii) Mealy. From the pattern point of viewa Mealy is identical
with a Grizzle but the White coloration is to a greater or
lesser extent replaced by Red. A Mealy (for Mealy
feathers see Pl. XX VI. figs. 5 & 6; the birds are figured
on Pl. XXV.) may therefore show Blue and Red, or Biue,
Red and White.

(iv) Blue (Pl. XXIII. fig. 1) is the colour of the Wild Rock
Pigeon, although in domestic breeds the rump is not
necessarily white. Ina Blue Chequer (Pl. XXIIT. fig. 3)
the light apical portions of the feathers are of the typical
blue colour, the rest of the feather and the general
appearance of the bird being very much darker. A very
dark Blue Chequer isalmost black, but this black is usually
dulland must not be confused with the glossy black (beetle-
black) characteristic of a pure black pigeon.

(v) Silver is a very pale blue with black bars and *‘ dun” flights.
In a Blue pigeon the flights are black (PI. XXIII. fig. 2).

(vi) Red. A Red pigeon is deep red all over including the flights
and tail. In a Red Chequer the dark portion of the
feathers are of the normal red and the light apical spots
white. The flights and tail are white.

It must be remembered that all these varieties show considerable
differences in shades of colour, and in the intensity and abundance
of the pattern markings. In this paper, however, we do not
propose to enter into these details, important as they are. Our
object for the present is to separate those characters which follow
apparently a Mendelian inheritance, from those whose mode
of inheritance is different. This paper therefore will only deal
with Mendelian inheritance, except to note in a few cases where
that inheritance is apparently subservient to other causes. None
the less these differences of shade and amount of pattern do
obviously follow a definite law of inheritance, and they are by no
means being disregarded by us, nor are we without hope of being
able at some future time to attempt some explanation of the laws
which govern their inheritance.

We may point out further that in the case of Mealies and
Grizzles (i.e. where normally there should be a small amount of
white), there is a great tendency to breed out in the course of a
few generations to practically pure white with only a few coloured
feathers. Our results, as far as they have gone, seem to show,
however, that these nearly pure white birds are still transmitting
to their offspring the colour which characterised their parents
and grandparents.

Blues and Silvers.

Blue is dominant to Silver.
A careful analysis of our results shows that Silver is in reality
a dilute Blue and that the colour factor in both is identical. The

604 MESSRS. J. LEWIS BONHOTE AND F. W. SMALLEY ON

allelomorph is therefore concentration and dilution (¢ and d) of
which the former is dominant.

Blue may therefore be represented as BBee or BBed, where B
represents the blue colour factor. Silver must. theretore be repre-
sented by BBdd.

Blues are said to almost always breed true, that is to say, that
they never throw any Silvers—this idea owes its origin, however,
to the fact that those Blues which are impure dominants (2. e.
containing the factor d) are of a poorer colour than the pure Blues
and have in consequence long been rejected by fanciers, so that a
race of pure dominants has thus been evolved *. Several of our
matings show that Blues or Grizzles that: are heterozygous as
regards cand d will throw Silvers.

The following are the results of our matings for these particular
characters, with the exception of Experiment 83, which is intro-
duced to show the reappearance of Silver from two heterozygous
Blues.

{Berenson bos IMGae alc ae lie ce:
| | Sales | ies eo foe ee EA DA Oe aR a as!
gs. |00/61 | Grizzle| BBed@g | — || 09/126 | Blue |BBedgg | — || 3B 1S.
14g. | 04/70 | Silver | BBdd — |}oa21 | Silver |BBdd | — | ANS
149. | 05/96 | BBdd — || 05/51 ‘! BBaa == |) ox
150. | 07/4 » |BBd@ | — [0656 | ,, | BBad =" Hh hs
151. | 05/96 so (eBiBad le Osan) Mines ea BBG: Zale os
152. | 03/5 i ee a BBddgg | 83 || 04/140 | Grizzle | BBee or ed | 157 All B or Equality
153. |0410 |, | BBdd_— | 140 {oat| » |BBecored| 77 || 3
154. | 04/14 | Grizzle | BBee or ed) 96 | 07/52 | Silver | BBdd | 151 4s se
185. | 08/1336 | Silver | BBdd | 148 | 08/1368 | Grizzle | BBee or ed| 137 | eS *
156. |03/5 | | BBaa | 88 |] 01/50 | Blue | _BBce | — | All Blue
fey 03sje- | 5, | BBdd 9) 83 (02/81 } 5. | BBee | — i
158. | 05/95 | Blue BBecored| 105 |/05/51 | Silver | BBdd | — ||, ,,
169. | 06/13 | ,, BBed 65 ||06/66 | ,, | BBad [es i Equality
160. | 03/5 | Silver BBdd 83 ||03/126 | Blue | BBcd = wpillind ss
Tete] OF . |. gl) PBBda > [i NOGereanerr | BBed: \ EB adie ds
162. | 04/52 Blue — BBed 156 || 06/56 | Silver | BBdd — | F
163, | 08/1335 Silver | BBdd 148 | 08/1887 Blue | BBed 154 | fs

6B28
S

Result. |

7B —
4B 28

When the Gametic Formula is in italies it implies that it is not definitely known. Only the

characters under immediate consideration are given in the Gametic Formula.

* See also remarks on chequering.

+ We must, however, point out here that in Mr. Staples-Browne’s experiments,
one or two blues, apparently heterozygous, behaved as homozygous and vice versa.

~ With regard to the numbering of Experiments—the numbers refer to our stud
book, and rather than number the experiments quoted successively in this paper, it
was thought best to have only one set of numbers so that references might at any
future time be easily made without risk of confusion.

COLOUR INHERITANCE IN PIGEONS, 605

From these matings we see that :—

(i) Silver to Silver (Exps. 148 to 151) gives nothing but Silver,
according to expectation.

(ii) Blue to Silver (Exps. 152 to 159) should give nothing but
3lues or Blues and Silvers according to whether the Blues
are heterozygous or not,and wesee that when our knowledge

of the gametic for mul of the parents was known for cer-
tain, the results were in exact accord with expectations, and
no results ant: vgonistic to possible expectations occurred.

(iii) Heterozygous Blues to Silvers (Exps. 160-163) should give

equality of Blues and Silvers. In the case of Exps. 160 and
161 no Silvers appeared, but the numbers bred were very
small. In the other experiments, exact equality was
reached, so that we may well assume that a continuation of
the other experiments would have led to the appearance
of some Silvers,

(iv) Heterozygous Blue x Heterozygous Blue should give Blues
and Silvers in proportion of 3:1, which was the exact

result of this mating (Exp. 83).

The results therefore show clearly that Silver is recessive to
Blue. The arguments showing that the difference between these
colours is one of concentration and dilution (rather than a differ-
ence of the colour factor itself), depend on the study of this
dilution factor, which, as it concerns the inheritance of other
colours and shades, not dealt with in this paper and at present
only imperfectly understood, is best deferred for the present. Its
discussion in no way affects the proof of the dominance of Blue
over Silver.

Chequering.

Chequering is dominant to pure colour. It is difficult to
realize therefore how it can have originated, since the typical
wild pigeon shows no such markings. At the present time,
however, in many parts of the country wild birds show
chequeri ing, but it seems more than likely that in these cases the
marking has been introduced by a cross with the domestic bird, as
most of the Wild Rocks in this country are now intermixed with
feral ones. Once this cross had been effected the chequering would
of course frequently show itself.

In direct contradiction to the foregoing remarks the following
fact is worth noting. Some years ago a pair of pure wild bir ds
was taken by one of the authors from a remote district in the
West of Ireland. All the wild pigeons seen (and there were no
tame ones within a radius of at least 30 miles) were purely typical
Blue Rocks. This pair bred in an aviary for five or six seasons,
producing only typical wild birds like themselves. Two seasons
ago (in 1909) an attempt was made to establish some of their
progeny as semi-wild birds and they were allowed to fly at liberty
from a dove-cot. One pair remained and reared several young, one
of which proved to be chequered !

Proc. Zoou. Soc.—1911, No. XLITIT. AB

606 MESSRS, J. LEWIS BONHOTE AND F. W. SMALLEY ON

Scientifically this result is of but little value, as the hen bird
may have made a chance mating with a chequered pigeon ; on the
other hand, the youngster was in every other respect (shape and
size) a ty pically wild bird. Unfortunately it met with an accident,
so that we have not been able to breed from it.

Unsatisfactory as this case is, it is obvious that the chequering
must in the first place have onginated from the Wild Rock
Pigeon, and the above is probably a good example of its arising
as a mutation.

A further explanation may possibly be found in the fact that the
chequering cannot show itself except in the presence of two shades
of colour, and that it may be present in many of the pure wild birds
but cannot show itself until the colour factor producing the two
shades is present, when the chequering will immediately appear.

This, however, in no way affects the main issue, namely that
when once a Chequer has been produced it is dominant to the pure
colour. The mating of two Chequers should therefore produce
Chequers or Chequers and Self-colour, according to whether the
birds are homozygous or heterozgyous for that character.

As in the case of the Blues, so in the case of the Chequers it
appears that fanciers, by continually selecting the best birds for
breeding, have unconsciously been selecting only homozygous birds,
with the result that only pure Chequer dominants are to be found
in certain strains. We have for instance accurate records of
57 matings (Exps. 1-49; 169-172; 175-178) of Chequer to
Chequer from which 229 young were produced, and all these
without exception were Chequers like their parents. It theretore
became essential to carry out further matings with birds that
were known to be heterozygous in order to test the dominance
of the Chequer character. This has been done in the matings
detailed below, the results of which, as will be seen, approximate
very closely to the Mendelian expectation.

f

9 Ext. Gametic | From | 3 | Ext. |Gametic | From Eanectdl Resale
App. Formula. Exp. App. |Formula.| Exp. tions. | aa
06/104. 4 ace } Xx | 173 ‘|| 06/82 | Blue. XX 122 | Equality 2Ch :2B
|
650. F Once i) Xx | 148-B |} 06/69 Grizzle.| xx | 158 x 3:2
lc G : ; | |
07/5 if Gate ; Xx 196 |kaS/e | Blue || xx | 196 be es
ye; | § Grizzle ; ail Weer | Bive a S| ‘
O7/a a Chequer Xx 196 || 08/d Chequer, Xx 196 | 3:1 | 2:2 |
07/3 E Gane ; Xx | 196 | 08/13! Grizzle.| xx | 196 |Equality) 2:0
Areal alate 27) 2 li pgjoo | Blue | 7 | ie ire
05/19 | 2 Chequer. Xx 197 1 08/22 | Chequer, Xx 197 3:1 10:1
ee ue ae || eps | ue = i .3%
09/533) } Chaguer s| Xx | 197 || 08/36 ‘Chequer, X* | 188) 3:1 | 3:34
08 24. | { Ghee : Xx 198 | 08/30 | Grizzle. NX 197 ||Equality}; 7:2

Chequering and its absence are represented by X and x.

* Matings for the purpose of testing the extracted recessives from Exps. 200 and
201 have been undertaken this year (1911), and prove the recessives to breed true.

COLOUR INHERITANCE IN PIGEONS. 607

It should perhaps be noted that although the individual matings
show, considering the small numbers, a very close approximation
to the expected results, yet at the same time the tendency to vary
from the anticipated results isall in one direction, viz. to a greater
number of Chequered birds. This is most marked in those
matings where equality was expected, for of the 28 birds bred,
19 were Chequers; in those cases where the-expectation was 3: 1,
21 birds were bred of which 15 were Chequers, which is approxi-
mately correct. It is possibly due as much to this tendency as
to the unconscious selection by breeders, that this character has
become perfectly true and stable in some strains. As pointed out in
the earlier portion of this paper, we are restricting our remarks for
the present to the consideration of the Mendelian inheritance of
certain characters, and that theory seems to fit in well with the
main lines of inheritance as borne out by the facts, None the less
it is equally evident that there are other factors at work, which
are able to modify to some extent the results anticipated by the
Mendelian hypothesis. In addition to these definite matings we
have also notes of 6 matings Chequer to pure colour (Exps. 164—
168, 173) which gave 19 birds all chequered, In this latter set
of matings most of the Chequered parents were birds used in or
bred from the Chequer to Chequer matings; and therefore this
adds further proof that all those birds were homozygous
dominants, as otherwise we should have expected some self-
coloured birds to appear as they did in Exps. 174-203 (p. 606).

It may be as well to mention here that although some of the
matings referred to in this paper were not undertaken with the
special purpose of bringing out the facts which they are used to
interpret, yet they have all been conducted by one of the authors
in person. Special matings have, however, been made in every
case to prove the inheritance of the characters discussed *.

Grizaling.

Grizzling is dominant to Chequering and hence also to pure
colour. It probably originated from the cross between Blue and
White, although such matings usually give splashed birds, owin
probably to the true Grizzle character, in which individual barbs
show both white and blue, being absent. Cases, however, are
known in which the cross between pure White and pure Blue
have produced Grizzles, and in these cases there is little doubt
that the Grizzle character must have been present in one or both
of the parents but was unable to show itself owing to the bird
containing only one colour. Once, however, the Grizzle has shown
itself, the White and Grizzle characters seem to combine together
and to have a common inheritance. Furthermore, as already
stated, Grizzles when bred together tend usually, but not
invariably, to show an increase of white in successive generations

* Mr. W. Bateson has stated (Mendel’s Principles of Heredity, p. 43 (1909)) that
chequering is dominant to its absence; on writing to him for a reference to the
source of his statement, he says that he had in mind some experiments of Mr. Staples-

Browne, which have, however, not been published.

608 MESSRS. J. LEWIS BONHOTE AND F. W. SMALLEY ON

till eventually some birds will be produced showing only one or
two coloured feathers. This matter, however, we shall not
discuss at present.

For our present purpose we may ignore the White character,
assuming that it is always present and inherited in common with
the Grizzle. The gametic formula, therefore, of a grizzled bird
will be BBee or ed GG or Gg, where B is the Blue colour,
e & d concentration or dilution (the combination dd producing
a bird known as a Silver Girizzle), and G & g the presence or
absence of Grizzle. Therefore from the crossing of two Grizzlies
we may either get all Grizzles or Blues and Grizzle es, according to
whether the birds are homozygous or heterozygous to the Grizzle
character. Under the term Blue we here for simplicity’s sake
include Silver.

Again, as in the case of the Chequers, the results from pedigrees
are apt to be misleading owing to the difficulty of distinguishing
the homozygous from the heterozygous birds ; nevertheless such
results entirely bear out our hypothesis, since, as regards the
colours produced, the results are quite in accordance with ex-

pectations.
Grizzle to Grizzle*,
|g | Bek [Gametic trom |g] Bet [Gametic | rom I papestation| Resut
74. 02/46] Griale |G@GorGg] — |/99/104 Grizzle| GG | — || All Gor3:1/3 Grizzles
75. |03/21 | % Gg 87 {98/88 ; iGGorGg] — || ,, » | 5 Grizzles
76. |00/21 a GGorGg) — ||63/7 ss KC ore == 5 | & Grizzles
77. |00/61 - Gg | 99/104) 4G — || ,, » | 7 Grizzles
78. |04/21 » |GGorGg) 76 04/58 é Ge oy ll. | 4G 2a
79. 104/23 . Ge 99 ||06/102) Gg | 153] 3:1 Me WS
80. |04/1 rev. “3 Ge 99 1105/6 | " Ge 109 |) 333 IL 1G —
| 81. jowie |, Gg 103 |o1/15 [SO 16Gror Gel == i AniGeor se fate anne
* 82. |06/88 55 GGorGg 78 ||04/58 x Ge 94 | AllGor3:1] — 1B

% As Theos aane the Gharacten dealt tans Is Shows nin the Cancun Woumula,

Looking through these matings, we see that when both Grizzles
and Blues should have appear ed the expected proportion ought to
have been 3:1, and the results give 10:5 or a rather large
excess of Blues.

Grizele to Blue.

These matings (Exps. 83-138, 152-155) may be divided into two
groups: (1) those in which the. gametic formule of both parents
were known ; (2) those in which, owing to the gamietic formula
of one par ent coe doubtful, two expectations were possible.

Certain of the Grizzles were known to be heterozygous: this was
the case in 19 matings, in which therefore Grizzles and Blues
should have appeared in approximately equal numbers. Alto-
gether 64 young were reared, 37 Grizzles and 27 Blues, thus
showing a slight excess of Grizzles.

; COLOUR INHERITANCE IN PIGIONS, 609

Grizzle to Blue.

| | i
0 IWxt. |Gametic From | Ext. |Gametie! From |, ee é
t App. |Formula., Exp. | 3 ADD. ‘Formula. SD: | Expectation. Result.
00/61 Grizzle |GGorGe| —_ || 02/126 nice ge | == “All Ge or Kqui Ting 4G 4713 |
01/27 Blue oo a= | 01/28 Grizzle |(GGorGe) — | 3 P 5G 1B)
O1/10 u | Bo — || 03/26 , |IG@GorGe — || ,, ss 1G —|
00/21 Grizzle \GGorGe| —_ || 02/35 Blue OM) ieee IML gi Po 1G 2B)
00/6 rev. | Blue go — || 02/24 Grizzle |GGorGg) — - F 2G 423
02/9 a | gor — || 01/24 » \IaGorGg) — || ,, 3A 2G 4B
02/37 | Grizale |GGorGg| — _ || 02/41 | Blue ge | — A, vs 1G 2B
02/46 (a) Pr GGorGg) — 21 i Pape Wo IN pe 3 4G —
02/46 (b)| 4, IGGorGg| — 43 - oor — | 5 - ==) 6B!)
03/31 Blue go 84 || 02/23 Grizzle |GGorGg, — | ,, i — 5B
01/27 ” £8 cae | O1/15 ” ” eta ” ” 4G 23 |
03/19 _ 0" 86 || O1/24 3 as — |, mm 2G 2B
03/11 iy ow 51 || 02/77 ss rile ileal ta | caer . 4G 1B
Ol /13 ” £28 baa (1/28 ” ” ee! ” ” 4G 1B
03/37 Grizzle GGorGg| —_ || 02/38 Blue ge — || 4 3 — 8B
03/20 fr Ge — || 02/22 55 oor — || Wauality 1G 2B
O1/11 Blue eo — || 03/9 Grizzle Gg | 84 | - 5G 1B
01/10 is ep — || 00/53 in GGorGe — |AllGgor Hquality) — 1B
NORTON Ns oo — || 98/96 iy oI oi i i 3G 3B
02/34: Guizzle |GGorGe| — _ || 02/26 Blue ge | — | Bt is — 1B
98/9 | - i — || 02/20 i | Popes Soll y a WG eus
00/6 Blue oor — || 03/63 Grizzle | Ge | 88 - Jquality 1G —
02/52 Bs | po — || 03/7 » |GGorGs, — | AllGgor Equality) 2G 4B
01/27 re oo — || 01/24 * IGG orGe) — - rf — 1B
02/46 (6) | Grizzle |GGorGe| —_ || 02/25 Blue on : r - 1G 5B
03/75 2 5. 74, || 03/51 “bi ge BBE Nt ie # 3G 2B
02/46 (a) - i. — || 03/23 os gor 53 Ci|| 2G 5B
04/31 | Blue go 59 || 03/9 Grizzle |} Ge | 84 " Wqu: lity 2G 1B
04/7 I ge 95 || 00/53 » MGGorGe| = All Gg or Equality) 1G 3B
04./4 is oo — || 01/28 55 IGG orGe) — if Ai 2G 2B
03/29 As oor 88 || 02/77 4 GG orGe) — os s 3G 3B
00/21 Grizazle |GGor Ge} —_ || 04/84 Blue | gg 157 os He 2G 3B
04/3 rev. | Blue oo 61 || 03/8 Grizale |GGorGg) — a ‘ Gielen
00/61 Grizzle Ge — || 04/12 Blue | gg 61 % 3 6G —
05/14 Blue gor 65 || 01/53 Grizzle GGorGg) — . 3G 5B
| 04/121 EY) ge 59 || 01/24 ” ” nei ” oy) Cr |
01/7 Grizazle |GGorGe| — || 02/22 Blue | gg — ii (3 2G 8B
02/46 (a) * — || 04/53 4 ge 61 58 is 12G —
03/75 BS x 74, || 04/48 fe geo a= a es 2G 4B
02/46 (b) e 3 — || 03/78 ‘) go 53 % i 1G 5B
O4/4rev. | Blue ge 61 || 05/6 Grizale | Gg 95 Kquality 3G 4B
04/93 re oo 61 || 01/53 » |GGorGe| — | AllGgor Equality!) 4G 1B
04/115 Griazle | Gg 104 || 04/120 Blue | ge 59 || Equality 3G 2B
04/133 i Gy 90 || 02/22 i og — || 216 —
04/52 Blue ge 156 | 04/10 rev.) Grizale GGorGg 77 | AllGe or "Equality! 4G —
05/40 Grizzle Ge 108 || 04/53 Bich mare 61 || quality 3G —
05/119 Blue ge 65 | 00/53 | Grizzle GGorGg — — AllGgor Equality) 1G 1B
02/46(a) | Grizale |GGorGg| —_ || )4/27 Blue ge | Ql \2q —
04/14 nt Gg 96 || 06/87 Fe fe fsafosy fh iL} ! ” quality 4G —
05/92 Blue oe 105 || 03/9 Grizale | Gg | 84 || i Gra lls
04/121 . Bo 59 || 04/58 aly Ger, |) Oa i [ete
05/96 AA oor 105 || 06/22 Ah a AG yea alteattalis) of Pe |) A Bay
04/13 Grizale | Ge 96 || 04/48 Blue | ge | — || cs |\4G 1B
06/57 | 6 Gg 118 || 06/63 55 | ge 123° || 5 — 1B
04/121 | Blue ge 59 || 04/58 | Grizzle | Ge 94 | ey 2G —
06/57 Grizzle Ge 118 || 05/43 Blue yp) Mila ‘ |} — 1B)
03/5 | Silver eo 83 || 04/140 Grizzle | Ge lal 3 Gaels
04/10 si ro 140 || 04/l0rey.| _,, GGorGeg) 77 | AllGgor Hquality) 7G — |
04/14 Grizale | Gg 96 || 01/52 Silver we Uy Kquality 2G 4B
08/1336 | Silver gg | 148 | 08/1368 | Griade | Gg | 137 3 | 2G 4B

610 MESSRS. J. LEWIS BONHOTE AND F. W. SMALLEY ON

In only 8 of the other matings did Blues fail to appear, and
we may therefore consider that when they did appear the
expectation should have been equality as before. Deducting
therefore these 8 matings which only produced Grizzles, we have
left 33 matings producing 156 young, 63 being Grizzles and
93 Blues. Since only one expectation is possible if both colours
appear, We are justified in uniting the figures from these two sets
of matings and treating them together. We find, therefore, that
in the total we have 52 matings of Grizzle to Blue, which produced
220 young of which 100 were Grizzles and 120 Blues; and this
proportion, although not exact, is not unreasonably far from the
expectation (Equality), and certainly seems to show a Mendelian
basis of inheritance.

If we digest the facts still further, we find that actual equality
was reached in only 19 eases, and this by including the odd
numbered broods where the deviation was not more than one.

We have, therefore,

Actual Equality reached in roughly 36°/, of the

matings.
Blues outnumbered the Grizzles by more than one in 20°/, ,,
Grizzles ss ., Blues = Es 12°(0 ss
Blues only, appeared in ilo) Meee
Grizzles only, : _ PAS Joment Ez

This, therefore, seems to show that while only a moderate
percentage of matings gave the exact Mendelian expectation, the
variation to one side or other of the mean is fairly evenly balanced,
with however a slight but unmistakable tendency towards an
overproduction of Blues: a tendency which was also shown in
the Grizzle to Grizzle matings.

Thus, as in the case of the Chequers where we found the
Mendelian proportions fairly well maintained, but with a distinct
tendency to an overproduction of Chequers, so also in the Grizzles
we see a similar tendency to an overpreduction of Blues.

There remains for consideration the 8 matings in which only
Grizzles appeared. Two of these may be at once dismissed as
only one bird was reared, so that we have no hint as to the
probable expectation. From the other 6 matings 25 birds were
reared, so that there is a reasonable probability of a Blue having
appeared were either of the parents heterozygous. Unfortunately
for the simplicity of this reasoning, we find on investigating the
matter more closely that although in 3 of these matings the
same hen was used and we might therefore presume her to be
homozygous, yet by her progeny in another mating (Exp. 109)
she proved herself to be heterozygous. This then leaves only 3
cases out of 60 in which the Grizzle parent might be homozygous.
So that again, just as we found in the Chequers and Blues an
overwhelming proportion of homozygous birds, which we attributed
to the unconscious selection of breeders, similarly in this case the

* This last calculation is of course exclusive of the 8 matings mentioned above,
in which the gametic formula of the Grizzle parent was doubtful.

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612 MESSRS. J. LEWIS BONILOTE AND F. W. SMALLEY ON

number of the heterozygous birds largely predominates, and this
is probably due to the fact that Grizzle x Blue is the commonest
form of mating used by breeders to produce Grizzles.

We have no records of matings with 2 extracted (Grizzle-bred)
Blues, but of Grizzle-bred Blues to Blue we have made 9 matings
(Exps. 139-147). These matings produced 28 birds all Blue in
accordance with the expectation.

Tn order to still further test our hypothesis we have carried out
the foregoing experiments (see Table, p. 611).

The result of a ser utiny of these matings is sufficient to prove
the Mendelian inheritance of the Grizzle “character. In some of
this last batch the expectation was well defined ; in others—owing
to the impossibility of distinguishing homozygous and hetero-
zygous birds—the expectation was open to one or two, or in some
cases three, interpretations. In those cases where the expectation
was all Grizzles or Grizzles and absence, in proportion of 3:1, and
only Grizzles were produced, we have concluded that one of the
parents, at least, was homozy gous—similarly, if any of the pro-
geny in those cases lacked the Grizzle, we have presumed that
the expected result should have been 38:1. On this basis we
have tested the results and we find :—

No. of
Expectation. Matings. Result.
AUG a 27 1ESy es ccteuys 7 39 Grizzles.
BE ea Aa aire Helin s ar eee 32 Grizzles, 11 absence of Grizzle.
ADH GHP aeaen ace ec nncne cds 3 9 Grizzles, 9 absence of Grizzle.
INOMGIZZlesm Byaceen tones 2 18 absence of Grizzle.
Higuelityror 3: bean. Sit. aie 5 Grizzles, 1 absence of Grizzle.

This last set of matings places therefore beyond doubt the
Mendelian inheritance of the Grizzle character.

In the summary given above we have, however, left ont
Exp. 180, in which with an expectation of 3:1 or all Grizzles,
14 Grizzles and one pure Blue were produced. At the present
moment, we can offer no reason for this considerable deviation
from the expected result.

Grizales and Chequers.

The Grizzle character is dominant to the Chequer, although,
in almost every case, the heterozygotes may be easily recognised.

To test this inheritance we have made the following matings
(see Table, p. 613).

These matings show fairly clearly the mode of inheritance,
but from the smallness of the numbers the proportions of the
different colours are not always in exact accordance with the
expectation. It may be further noted that although, from the
above reason, certain expected colours have as yet not occurred,
on the other hand no unexpected colours or combinations have
appeared, Thus we see that in Experiments 200 & 201 the
recessive Chequers: gave us, as expected, 14 Chequers to 4 Selfs,
and no Grizzles; in Experiments 202 & 205 the pure Grizzles
gave us, 11 Grizzles to 1 Self, and no Chequers. These two latter

6

613

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614 MESSRS. J. LEWIS BONHOTE AND F. W. SMALLEY ON

matings (Exps. 202 & 205) may be compared with Exps. 203 &
204 *, ‘matings of heterozygote Grizzles, where two Chequered birds
turned up, although the whole number bred (12 birds) was far
too small to give any hope of an approximation to the expectation
being attained, Many of the Grizzles in these matings are very
white, a factor which would tend to conceal the Chequer markings
if present. It is more than likely therefore, that owing to this,
some of these so-called Grizzles are in reality Grizzle-Chequers.
Taking the Grizzles and Grizzle-Chequers (of Exps. 203 & 204)
together, the expectation is 12 Grizzles and Grizzle-Chequers,

4 Chequers and 2 Self, and our result, 12 Grizzles and Grizzle-
Chequers and 2 Chequers, i is reasonably near the anticipation.
The expectation in Experiments 179, 193, 195, 196 & 197,
was an aqme of Grizzle-Chequers, Chequers, Grizzles and Selfs,
and the result 5 Grizzle-Chequers, 12 Chequers, 5 Grizzles, 8 Selfs,
the only real discrepancy here being the overproduction of
Chequers. This apparently merely confirms the conclusion come
to in an earlier part of this paper, p. 607, which showed that
there is apparently some factor which overrides the Mendelian
inheritance, and leads to an increase in the number of Chequers
produced.

Mealies.

A ‘Mealy’ may best be described as a Red Grizzled Pigeon
showing Blue. The general appearance may be seen by merercnee
to Plate XXV, and it should be noticed that it has the red bars
and white flights characteristic of some varieties of red pigeons,
When dealing with the Grizzle-character (G) we had to consider
its relation to White, and we came to the conclusion that the
White was not a colour factor complementary to Blue, but a
separate allelomorph; so that the real gametic formula of a
erizzled bird was made up of a compound allelomorph containing

three characters—blue, white, and grizzling (B, W,'G).) Bor

practical purposes, however, the W & G combine in their inherit-
ance, and thus in the cases we have been considering they have,
for convenience, been regarded as a single character (G).

In considering the inheritance of the Mealies, however, we
must again pause to consider whether the factor for Red is to be
treated as a colour factor, complementary therefore to Blue, or as
complementary to the factor for White, or yet again as a separate
allelomorph.

At first sight it would seem natural to consider it an alternative
colour to Blue, but if this be the case a certain number of Self
Reds should have e appeared in our matings. None, however, were
produced, though a certain number of Blues have been reared.

The same argument, though in a lesser degree, should hold
good if it had an inheritance of its own, and we are thus driven
to the conclusion that the Red is a complementary factor to the
White. We have also had certain aberrant results (not deait
with in this paper) which point to a curious connection between
these colours.

* This pair has produced both CG and pure Chequers this year (1911).

COLOUR INHERITANCE IN PIGEONS. 615

Finally, the study of the Mealy itself bears out this contention.
A dark ‘ Mealy’ differs from a dark ‘Grizzle’ in the fact that
the white portions of the latter are replaced by red and the
flights and tail instead of being black are white. In the Grizzles
we noted that there was a great tendency for the white to
increase till an almost pure white pigeon, showing only a few
coloured feathers, was produced. Matings of Mealy to Mealy
show an increase of the white, as is the case with Grizzles, till we
eventually get a white bird showing a few coloured (Blue or Red)
feathers. We must then come to the conclusion that in Mealies
it is the white that is replaced by red, and not the blue, and
therefore that a Mealy is a Grizzled bird in which the white is
wholly or partially replaced by red.

We have had.to go into this matter thoroughly as it offers
certain difficulties, which cannot be entirely cleared up till the
relationship of white and red have been further investigated ;
nevertheless the results of our experiments will offer no difficulty
if the red is considered as an alternative factor to the white.

In onr experiments with Mealies, one character has been
present in all the matings, namely Blue, or in its dilute form
Silver; we may therefore dismiss it from our calculations.

The only Chequer which appears (the one in fact by which the
- red colour was originally introduced) is shown in Exp. 179;
the only Grizzle bird from that mating (Mealy ¢ 54), whose
descendants form the large bulk of the Mealies, emphasizes the
truth of his inheritance, since no chequered bird has appeared in
spite of the large number bred.

The Grizzle character in the Mealy or Mealy bred birds we
have already dealt with (Exps. 180-189, p. 611). The question
therefore left us to consider in dealing with the Mealy inherit-
ance is the question of Red and White. Red is apparently
dominant to White, and in consequence a Mealy is dominant to a
Grizzle.

The following matings (see Table, p. 616) show the inheritance of
this character. It must be borne in mind that W & w are in this
case practically equivalent to G & g in the earlier part of this paper.
According to our present knowledge, when the Grizzle character
meets with either white or red they combine in their inheritance
to give either a Grizzle or a Mealy.

Taken as a whole, it will be seen that the results come remark-
ably near the expectation. In 5 of the 11 matings, in spite of
the small numbers, the results exactly bear out expectations ; and
in all the others, with the exception of Exp. 181, the results are
sufficiently near to leave little doubt that a continuance of the
mating would hare made them correct.

Taking the expectations and results together but omitting Exp.
181 we get :—

Expectation 2M.1G.1S. No.of Matings 4. Result 18. 10.9
a 2M 2G 0S © 5; Pe peed 4-8-0

in both of which sets there is a slight tendency to fewer

MESSRS. J. LEWIS BONHOTE AND F. W. SMALLEY ON

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COLOUR INHERITANCE IN PIGEONS, 617

Grizzles than one would expect. So that here again, although
the numbers are smatl, the Mendelian proportion seems to be
upset by a slight tendency in a definite direction.

The occurrence of a pure Blue (no. 2) in Exp. 181 is the only
instance throughout the whole of our matings in which a colour,
other than one which was expected, has appeared. Possibly the
male parent of this mating (1326), although unlike his brother
(1354) in external appearance, had in reality the same gametic
formula. Such an occurrence would not be without precedent, as
Mr. Staples-Browne instanced a homozygous Blue which showed
a certain amount of white, and this ought to have indicated a
heterozygous bird. The difference between these two birds (i. e.
presence of white) is of the same nature, and in support of this
suggestion the result of the mating—7, 2. 1—seems to be following
on the same lines as Exp. 182.

There is one other possible suggestion for this abnormal result,
and this is that the White or Red character may not have been
inherited with the Grizzle.

The full formula of the parents on this basis is :—

2 BBed Gg [Ww|— ¢ BBed Gg [RW].

Now if the Grizzle has been independently inherited, we might
get BBedgg [RW, WW, wR, Ww] as the formula of one of
the offspring, and the last character | RW ete.] might not in the
absence of G be able to show itself, and this would give us what
we got, namely a pure Blue. The result of the mating of this
self-coloured bird (Exp. 192) throws no light on this, for if, as is
oS possible, her mate was a homozygous Grizzle, no selfs would

uppear.

It must be understood that these are merely possible explan-
ations, of which we are inclined to favour the first, but at present
we have no definite proof in support of either.

For the rest, we claim that our hypothesis is so closely borne
out by the facts that 1t may be accepted till further work
confirms or disproves it: and until we are clearly able to
differentiate between those characters which follow the Law of
Mendel and those which are apparently governed by other laws.

In this paper we are only touching on the fringe of colour inheri-
tance in Pigeons, as there still remains the question of Black,
Dun, Red, Yellow, and White inheritance, on which we are at
present continuing our researches. Our work, however, empha-
sizes the fact that there are three important problems which the
Mendelian hypothesis fails to meet :—

The differences of shades in the same colour ;
(ii) The predominance of one sex in certain colours * ;
(i) The gradual increase of the white in Grizzles and Mealies
in successive generations ;
* This most interesting question has not been dealt with in the present paper,
as we have not yet fully nivestic ated the results ; but we may mention that a large
proportion of the White Grizzles are 2’s, and in the Light Mealies by far the

larger number are @’s; we have also bred a certain number of Cream Mealies, and
these have all been ?’s.

618 MESSRS, J. LEWIS BONHOTE AND F. W. SMALLEY ON

and in addition to these the apparently large predominance of
homozygous Chequers and heterozygous Grizzles.

These questions have been very much before us during the
whole of our experiments, as Indeed must be the case, for they
cannot fail to bring themselves to the notice of every practical
breeder.

At the present moment we do not consider it advisable to
bring forward any attempted answers to these questions. Much
more work yet remains to be done among the other colours, and
until we know more of their inheritance any attempt to solve
these problems would be premature. On the other hand, we are
not without hope that the further experiments in which we are
at present engaged may help to throw some light on these
perplexing problems.

v
Summary.

This paper may strike a reader as having for its main object
the confirmation of the Mendelian Laws; that, however, is by
no means the case. Realizing that many details of inheritance
did not entirely accord with the Mendelian theory, our object has
been to extract, so to speak, from the results obtained by mating
on Mendelian lines, that portion of them which clearly shows the
Mendelian inheritance. ‘The residue must, we submit, have been
brought about by some law or series of laws, which overrides and
modifies (externally at all events) the expected Mendelian results.
By means of this extraction we are able to see the effect of that
law or laws untrammeled by the effects of inheritance as ruled
by the Law of Mendel, and thus we have been brought slightly
nearer to an understanding of them.

Now this paper deals with the Mendelian inheritance of the
characters considered ; consequently all details of shades of colour,
predominance of one sex in certain colours, and several other
similar matters have been entirely omitted.

On the other hand, certain points such as the superabundance
of Chequers in our Chequer matings and of Blues in the Grizzle
and Blue matings, have had to be brought forward and the results
are, we hope, sufficiently conclusive to prove to our readers that
although the characters dealt with follow in the main on the lines
of the Mendelian inheritance, yet it is equally certain that there
is another factor which is able to dominate and influence that
inheritance.

So far asthe matter has been dealt with in this paper there is
no evidence to show that the gametic inheritance has been
affected. Except in one doubtful case (Exp. 181) we have not in
the course of all our matings bred a single bird that was not a
possible result of the mating under the strictest expectation of
the Mendelian theory. What, however, we have been able to
show is, that in certain cases a consistent deviation from the
expected proportions occurs. It would thus seem at first sight as

COLOUR INHERITANCE 1N PIGEONS. 619

if some factor exists which has the power to influence but not to
alter the gametic inheritance.

The Mendelian conclusions reached in this paper may be briefly
summed up as follows :—

(1) Silver is dilute Blue.

(2) Blue is dominant to Silver.

(3) Chequering is dominant to its absence (7. e. a Self-colour),

(4) Grizzling is dominant to its absence (i. ¢. a Self-colour).

(5) Grizzling is dominant to Chequering ; the impure dominants
may however sometimes be easily distinguished.

(6) A Mealy is a Grizzled bird with the White wholly or
partially replaced by Red.

(7) Red in a Mealy is apparently dominant to White, and
hence a Mealy is dominant to a Grizzle.

(8) White and Grizzling when they have met combine together
and have a common inheritance.

(9) Red combines with Grizzling in the same way as does White.

EXPLANATION OF THE PLATES.

Prare XXIII.
. Blue Pigeon.
. Silver Pigeon.
. Chequered Pigeon.

ea a

PLATE XXIV.

. Dark Grizzled Pigeon.

. White Grizzled Pigeon.

. Grizzle and Chequer (Almond Grizzle) Pizeon—in nest feathering,
showing adult feathers appearing on the wing-coverts. (Note chequering
on the adult wing-coverts.) ;

wrt

PLATE XXV.

. Dark Mealy. Pigeon.
. Light Mealy Pigeon.
- White Mealy Pigeon. (Tricolor.)

wre

Pratt XXVI.
Feathers showing details of pattern-markings.

. Chequered feather.

. Dark Grizzled feather.

. Grizzle-Chequer feather, adult plumage, as shown in dark wing-coverts,
Pl. XXIV. fig. 3.

. White Grizzled feather.

- Light Mealy feather. (Note white. red, and blue in some barbs.)

. Dark Mealy feather. (Note absence of white.)

Fig.

On Whore

DR. R. T. LEIPER ON NEMATODE PARASITES.

EXHIBITIONS AND NOTICES.
April 4, 1911.

Dr. Henry Woopwarp, F.R.S., Vice-President,
in the Chair.

Sir E. Ray Lanxester, K.C.B., F.R.S., F.Z.S., exhibited a
special Supplement of the ‘ Field’ newspaper dealing with the
British non-migratory Trout, and called attention to this new
medium for the publication of scientific observations requiring
illustrations.

Dr. R. T. Lererr, F.Z.8., gave a demonstration of Nematode
parasites obtained from animals in the Zoological Gardens during
the vear ending November 1910.

The collection contained a number of new forms, of which a
systematic account will be published later. Among the more
interesting of the known forms were fictularia plagiostoma
from a Palm-Civet, a number of species of Polydelphis from
various Pythons, Dicheilonema horrida from the South American
Ostrich, and Dictyocaulus filaria from the lungs of Sheep.

It was noticed that whereas intestinal parasites were almost
wholly collected from animals that had not lived in the Gardens
for more than six months, those of which the normal habitat and
food were the internal tissues of the host occurred in animals
that had been confined in the Gardens for several years. Thus,
an undescribed /ilaria was found in a Lemur after four years’,
and Filaria australis in a Wallaby after two and a half years’
captivity.

In all these cases the number of parasites obtained was small,
and could have had little or no effect upon the health of the
host. There was a remarkable preponderance of female forms.

From these observations it appeared that the change of food
and general conditions obtaining in the Gardens were unfavourable
to the continued existence of the intestinal parasites an animal
may harbour on its admission. The number of cases of auto-
and re-infection during captivity was strikingly small, and bore
testimony to the cleanly surroundings in which the animals were
kept. In four cases only was there evidence of the occurrence
of accumulative infection in the Gardens :—

1. A number of Giant Toads died from lung infection with
Rhabdias bufonis.

2. The Wolves appeared to be heavily infected with Ascaris
canis.

3. A Sheep died from pneumonic condition resulting from
an intense infection with Dictyocaulus filaria.

4. The Tortoises had Oxyuriasis.

Tn all these cases repeated infection undoubtedly had followed

ON A NEWLY BORN CUB OF THE MASKED PALM-CIVET. 621

from contamination of food and drink with feces containing
eggs of the parasite. ‘The infection could be eliminated by steam
sterilisation of the cages, or still more easily by changing the
species of animal living in the particular paddocks or cages, for
Helminthes were often peculiarly selective as regards their hosts,
and those flourishing in one animal sometimes found it impossible
to continue their life even in closely allied forms.

Mr. R. I. Pocock, F.R.S., F.LS., F.Z.S., Superintendent of
the Gardens, exhibited the newly born young of the Masked
Palm-Civet (Puradoxurus larvatus), which had been born in the
Gardens from a pair from Szechuen, presented to the Society by
Mr. Thurlow Lay, and remarked that, although the specimen had
died soon after birth, two other individuals composing the litter
were alive and likely to do well. This was the first occasion on
which the species had bred in the Gardens. The coloration of the

Text-fig. 147.

Inner aspect of abnormal left fore-leg of a newly born Masked Palm-Civet
Paradoxurus larvatus.
p, pad; s, strip of naked skin; ¢, claw.

young resembled in a general way that of the adult, but was of a
more generalised type, the black and white pattern of the head
being less emphasized and the general colour of the body greyer
with less yellow; the greater part of the tail and the lower portion
of the limbs were sooty grey, the throat, chest, axille, belly, and
the inside of the thighs being white. Of special interest was the
presence of a pair of ill-defined dark stripes on the back and of
very indistinct traces of pattern on the sides of the body. The

Proc. Zoou. Soc.—1911, No, XLIV. 44

622 LAND TORTOISES IN THE SEYCHELLES.

head, which had the eyes and ears closed, was shaped very much
like that of a wolf-pup. The tail was short-haired and tapering
and as long as the body from the fore part of the shoulders
backwards.

Special attention was drawn to a peculiar abnormality of the
left fore-leg (text-fig. 147). The humerus appeared to be of
normal length, but the lower arm was quite short, and there was
no distinct elbow-joint ; the paw, although freely articulated at
the wrist, was axially rotated outwards so that its plantar surface
looked inwards. It was furnished with a single large pad repre-
senting the large pad of the normal foot, and was armed with a
terminal claw, a thin strip of naked skin passing from the claw to
the pad.

[Note added July 17th, 1911.—The two specimens of the litter
that survived grew with great rapidity as compared with dogs and
cats, and almost equalled the size of their parents when three
months old. Their eyes, however, opened, as in the former
animals, about the ninth day from birth. |

Land Tortoises in the Seychelles.

Tue Secrerary read the following dispatch from the Governor
of the Seychelles, a copy of which had been kindly sent him by
the Secretary of State for the Colonies, for communication to the
Society.

Government House,
Seychelles,

1st June, 1910.
My Lorp,

In view of enquiries made from time to time regarding
the conservation of the breed of land tortoises of the islands in
the Indian Ocean, I have the honour to append some notes
condensed from the entries in the stud-book of the herd at
Government House, Mahé. This book was opened by me soon
after my arrival here in 1904, and contains records of the annual
measurements of the specimens under my observation and of
their habits.

2. In June 1904 I found a herd of 42 adult land tortoises, and
17 young ones hatched out in 1902 and 1903; these were duly
marked, numbered and measured, and the particulars entered up
in a new stud-book. The bulk of the herd had been purchased
for the Government in 1892 from the late Mr. Nageon de!’ Estang
of Val des Prés, a proprietor of ancient family in the district of
Anse Aux Pins, Mahé. The animals were then transferred to
Curieuse Island, the property of the Crown, and were brought
back to Government House in 1902.

3. In addition to this herd, there are two large males : No. 1.
‘“Gordon,” presented by the late General Gordon (of Khartoum)
when he was stationed in Seychelles in 1881 after his transfer
from the Cape; this is the largest land tortoise in Seychelles, and
measures over the surface of the carapace 4’ 94". and 4’ 8". The

LAND TORTOISES IN THE SEYCHELLES. 623

plastron measures 3’ 9x 2'9". This is undoubtedly Testudo
elephantina according to Dr. Giinther’s monograph. The next
largest, No. 2. “Spurs,” was presented by Mr. Spurs, now of
Kuropa Island, a French possession in the Mozambique Channel :
I am indebted for many of my most curious notes on the habits
of all varieties of sea turtles and land tortoises to this gentleman,
who is an educated man trained in habits of observation and has
spent all his life among the islands of the Indian Ocean. The
present dimensions of ‘‘Spurs” are 4’ 5” x 4’ 4” on the carapace
and 3' 7" x 2'5'' on the plastron: he has grown slowly in breadth
since 1904 but not in length. This remarkably fine specimen (the
finest which Mr. Spurs has seen) is not of the elephantina variety,
and resembles the Testudo daudinii of Dr. Giinther’s monograph.

These are probably the finest specimens living of their race. My
recollection is that the Testudo elephantina which died at Colombo
in 1900, of a recorded age of 155 years, was of larger dimensions.
The large specimen at St. Helena was measured by Admiral Sir
J. Durnford in 1907 as 4’ 6” “ fore and aft,” but it is not certain
whether this measurement included only the shell of the back.

4. The adult females, which are readily distinguishable in
shape, are smaller than the males. The largest specimen in the
collection (No. 5) reached its present dimensions—3’ 8” x 3’ 10"
along the carapace and 2’ 8” x 2' 1” along the plastron—in 1906,
and has not grown during the last three years. Several others
have rather smaller dimensions and have not grown since 1904.

The breeding season extends from January to April: the
females carry their eggs for about 10 weeks and Jay them in holes
dug out by their hind legs and then covered over. The eggs in
each nest vary in number from 9 to 25 and are white, round, and
of the size of a lawn tennis ball. There may be two nests made
annually by one female. Sea turtles lay a much greater number
of eggs, e. g., the green turtle 250 eges at a time, and the hawks-
bill turtle 100 to 150.

5. The young hatch out in about 120 to 130 days and work
their own way out of the ground. At the Government House
“pare aux tortues” about half of the eggs are unfertile ; but in
some years of drought, very few young ones appear, being unable
(probably) to work their way up. They grow fast if well fed, and
at four years old measure 1’ 6" to 1’ 10” in length and breadth of
carapace. It is said that they attain full growth in 25 years. It
is a local custom to mark off a young one at any birth in the
family and to eat it at the child’s wedding day. The meat is
palatable and the liver is held to be a delicacy.

6. The number of young ones secured from 1904 to 1909 was
168; they are liable to be destroyed by rats before their shells
harden. Ina wild state at Aldabra practically all the young are
destroyed by floventins (cranes), rats, and wild cats.

7. Through the courtesy of Admiral Sir John Durnford and
Captain Dumas, R.N., six specimens from this herd have been
presented to various institutions, e. g., Groot Schnur, Pretoria, and
the Zoological Society of London.

44*

624 LAND TORTOISES IN THE SEYCHELLES.

8. Owing to the difficulty in providing food for the increasing
herd in the enclosures at Government House, I have drafted off—
in March and May 1910—4 adult males and 18 adult females
and 27 young ones of the “ récoltes” in 1903, 1904 and 1905 to
Long Island, a Crown property used as a quarantine station,
where they are placed under the charge of the Guardians and
where there is an ample supply of food.

9. “ Gordon ” shows likes and dislikes and is rather combative,
having successfully bitten some visitors who presumed on his
apparent lethargy, but generally the land tortoise shows little
intelligence.

10. There have been no deaths among the adults during six years
in the enclosures at Government House, but one male has been
killed by a fall at Long Island, where they have shown them-
selves to be capable of swimming. ‘Theremains of tortoises found
in the pits in the coral formation of islands in the Aldabra Group
points to the falling into pits as one of the principal causes of
death. ‘They live apparently to an extreme old age—probably for
200 years. No plan will effectively prevent the final extinction of
these curious survivals in a wild state in their natural habitats.
The archives of Seychelles, Vol. i., published in 1909, are full of
references to their size and number in Mahé and Praslin, where
they were speedily destroyed by the early settlers. But their
future existence is guaranteed by the fact that they breed in
captivity and that several large herds besides that at Government
House are kept and well cared for. It isa guarantee for their
being taken care of that there isa sale for living specimens for
zoological collections.

11. The best book in English on the subject is a monograph
entitled “‘ Gigantic Land Tortoises” by Dr. Giinther, published for
the British Museum about 1878. The names of the best known
varieties of the larger tortoises and turtles are as follows :—

Of Aldabra: Testudo elephantina.

Of Galapagos: Testudo nigra.

Of Greece: Testudo greeca.

Box Tortoise of Madagascar: Pyxis arachnoides.

Box Tortoise of North America: Ci%studo carolina (Brer

tarapin).

Lettered Tortoise of North America: Hmys sculpta.

Green Turtle: Chelone mydas. (The edible variety much

consumed in Seychelles.)

Loggerhead Turtle: Thalassochelys caretta. (1 do not know

this species by sight.)

Hawksbill Turtle: Chelone imbricata. (The tortoise-shell

variety.)
I have, ete.,

Tre Rigur HonovuRABLE Sioned W.E r
Tur SECRETARY OF STATE Pouginen) = 1 DASTDSOW,

FOR THE COLONIES. Governor.

DR. H. B. FANTHAM AND MISS A. PORTER ON BEE-DISEASE. 625

A Bee-disease due to a Protozoal Parasite (Nosema apis).

Dr. H. B. Fanruam, F.Z.S., and Miss Anniz Porter, D.Sc.,
exhibited some diseased bees and combs infected with a minute
pathogenic protozoal parasite, apparently the same as Vosema
apis found by Zander in diseased bees in Bavaria. Microscopic
preparations and drawings of the parasite, Vosema apis, were also
shown, as well as healthy bees and combs in contrast. The
material exhibited was obtained from Cambridgeshire and
Hertfordshire in. March, 1911. Some of the infected combs
were brown in colour instead of the normal yellow (combs of the
same age being compared), while the infected bees suffered from
a sort of dry dysentery which rapidly proved fatal.

The pathogenic agent of this dry dysentery, Vosema apis,
formed thousands of minute spores which fouled the hive, while
infection was probably spread to new hives by hungry, weakly
bees attempting to enter healthy hives. The spores, about 2 to
3p by 4 to 6u, were the resistant and cross-infective stages of the
Protozoon. The parasite Vosema apis was closely allied to that
of pébrine, the silkworm disease due to Vosema bombycis.

The trophozoite and pansporoblast stages of the Vosema apis
had been observed in the gut-epithelium of the bee. Some spores
with polar filaments extruded had also been found. It was very
probable that the young, growing and multiplicative stages of the
parasite were capable of killing the bees before the formation of
spores had been attained, for dead bees were often found in which
only young stages of the parasite could be detected, occurring
especially in the chyle-stomach and intestine. Like V. bombycis,
the bee-parasite was possibly capable of hereditary infection, as
infected bee-larve and a dead infected queen had been found
and examined. Maassen had recently found infected drones in
Germany, but the infection in drones was stated to be limited to
the intestine.

That Nosema apis was fatal to bees and allied Hymenoptera
had been shown by the exhibitors by feeding healthy hive-bees,
mason-bees, and wasps with honey infected with Vosema spores ;
also by placing hive-bees dead of the disease among healthy hive-
and mason-bees and wasps, and by direct contamination of healthy
bees with infected fecal matter. In each case the insects ex-
perimented upon succumbed to the effects of Nosema apis.
In Nature the method of infection is probably contaminative,
healthy bees becoming infected by swallowing the spores of the
parasite.

It should be noted that the virulence of the parasite appeared
to vary in bees at different times of the year and in different
localities. Bad seasons are usually followed by increase of disease.
Some bees became chronics, forming reservoirs of spores and so
acting as parasite-carriers.

The only certain destructive agent of the Microsporidian
spores was fire, and all infected bees and hives, and any débris
therefrom should be most carefully burned.

626 MR. F. E. BEDDARD ON

In the opinion of the exhibitors, the Microsporidian parasite,
Nosema apis, had been responsible for much of the bee-disease
recorded in this country since 1906, especially in 1906, 1907, and
1911. The exhibitors first noticed the parasite in 1906 in
diseased bees obtained from the Isle of Wight ; its full significance
was grasped in 1907, but owing to the difficulty of obtaining
material the exhibitors’ results were not published. As much
attention was now being directed to ‘“ bee-disease,” the exhibitors
briefly recorded their observations. It was not asserted that
microsporidiosis was the only disease of bees current in Great
Britain at present, as Dr. Malden had investigated a bacillary
infection in bees, the parasite being called Bacillus pestiformis
apis. ‘¢ Foul brood” also was a well-known and separate disease.

Microsporidiosis (due to Mosema apis) had probably been
introduced from the Continent into British apiaries.

Other parasites found in bees—chiefly in the gut—by the
exhibitors were various species of Gregarines, a Flagellate
apparently belonging to the genus Crithidia, a new Ameeba
(Entameba apis) very like Hntameba coli of the human intestine,
a Spirochete, and various Fungi.

PAPERS.

29. Contributions to the Anatomy and Systematic Arrange-
ment of the Cestoidea. By Frank H. Bepparp, M.A.,
E.R.S., F.Z.8., Prosector to the Society.

[Received and Read April 4th, 1911.)

(Text-figures 148-159.)

I. On some MAMMALIAN CESYOIDEA.

I propose to communicate to the Society from time to time
reports upon the species of Cestoidea which have been collected,
and are being at present collected, from animals which have
died in the Society’s Gardens. The collection in my hands is the
result of nearly two years’ examination of (necessarily) a great
number of animals, but does not contain as yet a very large
number of species, either of known forms or of those which
I believe to be undescribed. Tapeworms are by no means so
common as other parasitic worms, particularly Nematodes, which
are most abundant among the animals in the Gardens. Of
the forms which I have identified as belonging to well-known
species, I propose at some date to give a complete list, which will
be useful, not only as indicating the species which are most
abundant in the captive animals, but as extending the range of
hosts. At present I could hardly give a long enough list to

SOME MAMMALIAN TAPEWORMS. 627

warrant publication in the ‘ Proceedings. The new forms,
however, [ intend to describe, a few at a time, as soon as I
can give a sufficiently comprehensive account of their structure
to make such publication useful. The present communication
contains a fairly full anatomical account of three new species, of

Text-fig. 148.

Oochoristica sp., X 2.

two of which I have abundant examples for study. Of the third,
IT have been able to study two specimens only. But I have
been fortunate enough to fill up most of the descriptions which
are chiefly required to determine its systematic position with
accuracy.

On a Species of Oochoristica.

I vefer a number of individuals from the Lesser Anteater,
Tamandua tetradactyla, to this genus, for reasons which will be
apparent in the course of the following account of this Tapeworm,
and which are summed up at the end of the description in con-
sidering its affinities. I believe, however, that the present species
is to be regarded as a new one, though it undoubtedly comes very
near to Oochoristica wageneri, described by von Janicki from the
same Edentate four or five years since*. The reasons for this

* y. Janicki, “Studien an Satigetiercestoden,” Zeitschr. f. wiss. Zool. lxxxi.
1906, p. 5.

628 MR. F. E. BEDDARD ON

opinion will also be given later, after I have detailed the
characteristics of these worms.

As is the case with other species of Oochoristica, the scolex of
the present species is entirely unarmed and without rostellum.
I have studied this and other regions of the body not only by
inspection with a lens and the microscope of specimens in spirit
mounted entire in Canada balsam, but also by means of transverse
and longitudinal sections. I find that the rostellum is not
represented by the smallest vestige that can be recognised. The
anterior end of the head or scolex is simply slightly and uni-
formly convex, the convexity representing a large circle and, there-
fore, not to be described asa papilla. The suckers appear to stand
out rather from the scolex, and are somewhat long in shape with
a narrow and slit-like orifice.

The general aspect of the scolex itself does not recall that of
such other members of the genus Oochoristica as have been

Text-fig. 149.

Scolex and a few anterior proglottides of Oochoristica sp.

figured, excepting in its tetragonal form in section. It is, more-
over, rather unlike that of other Tetracotylea. When viewed
superficially with a lens, or under a low power of the microscope,

SOME MAMMALIAN TAPEWORMS. 629

the scolex appears to be of an oval contour, passing thus gradually
into a neck, which is at first narrow and later widens out. There
is nothing abnormal about the shape of such a scolex. A close
examination, however, shows that such a description would be
quite incorrect. For the scolex itself is in reality almost semi-
circular in outline, and its outlines are carried on on either side
by the neck of the worm, which is, to begin with, rather wider
even than the thickest part of the scolex. This is clearly to be
seen in text-fig. 149. It seems to be imbedded in the first seg-
ments of the strobila, like an egg in an egg-cup. Thus the true
scolex is very short. The widening of the strobila, for a space of
some five or six proglottides, suggests the commencing formation
of a ‘‘ pseudoscolex,” such as is further developed in, for example,
Thysanocephalum crispwm *.

The first steps in the formation of a pseudoscolex might well be
imagined to occur in some such way as is shown in the anterior
proglottides of the present species. There is, however, no further
indication of the formation of a pseudoscolex. The anterior pro-
glottides are precisely like those which follow, save in their
greater diameter. The figure also suggests that the scolex is
retractile within the first part of the strobila. In a sense this
is the case. Longitudinal sections through the scolex and the
commencing strobila show a bulging of the latter. There is a
plain demarcation between scolex and strobila, and not much
curving back of the anterior proglottides when thus viewed.
That the scolex itself may be extended to a greater length is
possible enough. And yet in seven or eight examples which I
examined there was an identity of structure in this and other
particulars. It is to be noted, therefore, that this species,
unlike Oochoristica tetragonocephala, has no neck, as, indeed,
was asserted of the latter by Diesing, but denied later by Liihe
and Janicki.

In examining the scolex mounted entire, after clearing with oil
of cloves or in alcohol previous to mounting, the suckers are by
no means so conspicuous as in other Tapeworms. The slit-lke
orifice can, indeed, be detected, but it is rather shrouded and dim.
The reason for this is apparent when the scolex is studied by
means of transverse sections, in which the relations to the head
ave shown. It will be there seen that, except for the small
orifice of the sucker, that organ is not by any means entirely
external in situation ; the sucker is, in fact, covered by a thickish
layer of the ordinary parenchyma of the body. It is thus imbedded
in the head, and represents a still further pronounced sessile
condition of the sucker, which (inter alia) distinguishes the
Tetracotylea from the Tetraphyllidia. At the same time, the
sucker retains its cup-like form, the two edges although in

* Cf. Benham in pt. iv. of ‘A Treatise on Zoology,’ ed. Sir E. Ray Lankester,
p. 121, fig. xvi. 2.

+ Unless, indeed, the anterior bulging part of the body of the worm is the neck.
But as it shows transverse furrows (text-fig. 149) I do not think that this is the case.

630 MR. F. E. BEDDARD ON

contact not uniting, thus leaving aslit-like orifice, which is dimly
seen when the scolex is viewed in its entirety.

The general aspect of the body is like that of other Tapeworms,
in that the anterior region is much narrower than the posterior
region, where the segments are in every way larger. In the
present worm this distinction is very marked, and the posterior
segments are rather more than oval in section, and approach
a circular contour. At the same time, this region of the body is
much stronger and thicker in every way than the anterior region,
so that there is a tendency for this part to become broken off.
There seems to be a rather sudden transition between the anterior
and the posterior regions. The posterior region of the body, in
fact, has quite the contour of an Earthworm, and entirely lacks the
flattened appearance of a typical Cestode. The cause of this
differentiation is doubtless to be sought in the swelling caused by
the dissemination of the ripe eggs through the medullary layer.
It is, in fact, in the posterior region that the uteri of the pro-
glottides have entirely disappeared, and are replaced by eggs
scattered singly through the parenchyma of the middle layer of
the body.

This state of affairs was, however, only to be noted as existing
in a marked fashion in the larger and, therefore, presumably more
fully mature examples of the worms. Fresh specimens reached
about 60 mm. in length, though in none did the breadth of the
body exceed 2 mm. Indeed, 1-1-5 was the prevalent diameter.
In smaller worms of 30-40 mm. length, the posterior pro-
glottides were by no means so swollen and remained comparatively
flat. So far as can be seen from an examination of the dozen or
so of specimens of this worm which I have in my possession, there
is no detaching of proglottides. In the first place, I always found
the terminal proglottid in its place, and to be distinguished from
the others by its oval posterior boundary interrupted by a large
orifice (apparently of the water vascular canals) highly suggestive
of a proctodceal invagination.

Secondly, the posterior set of segments were not materially
different in their form from those preceding them. In those cases
where proglottides are shed singly, it is common for them to
assume an oval form at the end of the body, and at least to be
a good deal constricted at their junctions with each other. There
is, commonly, also a very considerable increase of length of the
individual proglottides, which favours an easy rupture of their
connection. In none of these features do the terminal proglottides
of this Oochoristica agree with those of the genera which con-
stantly shed thei proglottides singly. Indeed, I have observed
that the five or six terminal proglottides are occasionally even
shorter than those which immediately lie in front of them. These
facts, coupled with the absence of a “neck” where new proglottides
ave formed, incline one to the belief that the growth of a scolex
is limited, and that the whole posterior region dies and liberates
the eggs. But of this I have no direct evidence. The relative

SOME MAMMALIAN TAPEWORMS. 631

length and-breadth of the proglottides differ in different regions
of the body and in individuals. A small example (30 mm. in
length) mounted in two pieces in Canada balsam showed the
following characters of strobilisation. The diameter of the wider
posterior region of the body was not more than three times that
of the narrowest part of the neck. 1 made out in all 167 strobila,
and there were not, I am convinced, many more. The first 42
were very short, being very much broader than long. After this
point the diameter of the worm became greater, and the segments,
therefore, larger; they appeared to be longer in proportion to
their breadth; but the first 31 of the series were, I believe,
not so. From this point, however, to the end of the body the
length of the segments was proportionately a little greater. But
throughout the series the actual breadth was really considerably
greater than the length. In a second specimen, of which the
head end only was mounted, the state of strobilisation presented
quite a different appearance. The first 45 segments or so were
quite as in the last specimen; but after this point the length
grew very rapidly in proportion to the breadth, and in a few seg-
ments no measurements were needed to assist the eye in seeing
that the proglottides were very much longer than broad. A care-
ful examination of these elongated segments showed that the
broader hind margin which overlapped the narrower anterior
section of the ensuing segment was regularly crumpled, showing
a series of regularly arranged projections, and in the middle of the
segment these were represented by alternate furrows and elevations.
This looks very much like the effect of a muscular pulling out
of a segment from end to end. But in any case this variability
in the proportions of individual proglottides shows that it is
a difficult matter to define Tapeworms very accurately by the
proportions of length and breadth of the proglottides in different
regions. The instances given would appear to be those of pro-
glottides which had been unduly lengthened through pulling
longwise. But I have observed instances of the contrary. In some
proglottides from the middle of the body there was a marked
transverse wrinkling, a kind of division of the segment into
annuli, which shows from a different point of view how difficult
it is to rely upon the relative dimensions of proglottides as aids to
the definition of a species or of a genus, as the case may be. It
is certainly quite unsafe in the case of this Oochoristica.

Before proceeding to the consideration of the internal organs,
there is one other external feature that requires description,
namely, the orifices of the generative ducts. ‘These are alternate,
but not regularly so. The openings are, however, as a rule, pre-
ponderatingly upon one side of the body. Thus in a portion of
one worm I found two pores consecutively upon one side of the
body. The next was on the opposite side; the five following
reverted to the first state, and the last examined showed again an
alternation. In another specimen, from another worm, there was
one pore on one side, followed by three consecutive pores on the

632 MR. F. E. BEDDARD ON

other side; then came two on the opposite side to these, and then
two on the opposite side to the two last described. These instances
will be sufficient to illustrate the general facts. It is quite
possible that the proglottid which first shows these openings may
vary from individual to individual. In any case a careful
examination of a small individual mounted in two pieces upon
a slide, and whose strobilisation has been already described above,
showed no generative ducts in front of proglottid 113 or so. It
will be remembered that this worm had a limited number of
proglottides altogether, there being not many, if any, more than
167. In this particular case it was not difficult to ascertain
accurately the point at which the ducts began. For, where
present, they were extremely conspicuous through staining, and
the cells composing the conjoined ducts formed a rather thick mass.
And I am, therefore, sure that they did not exist, except perhaps
in small rudiments, before the 113th proglottid. It was easy to
ascertain that the gonads themselves—at any rate, the ovaries
—existed anterior to this segment. Furthermore, these facts
ave still further supported by the condition of a second specimen
which had been cut into longitudinal sections up to about the same
point of the body. Here, too, the generative ducts did not exist
much anteriorly to the 113th segment—if, indeed, at all anteriorly.
The actual pores were only clearly visible upon more mature pro-
glottides, and were quite conspicuous round orifices near to the
anterior border of the proglottid.

The excretory vessels would seem from the figures of Janicki
to be very characteristic of the genus Oochoristica, for in the species
figured by him there are additional lateral vessels varying
according to the species. I have found the same thing in the
species of Oochoristica which I describe in the present paper.
The conditions, however, seem to vary somewhat in different
regions of the body. Anteriorly there are eight, longitudinally
running, which in the very anterior and therefore very thin
segments are almost in the same plane, and therefore can be seen
ina single longitudinal section. Further back the eight tubes are
differently arranged. There are two nearer to the middle line
and dorsal, according to Janicki*, and two ventral more widely
separated. The two lateral tubes on each side are less markedly
dorsal and ventral respectively. In posterior segments I could
see only six longitudinal vessels, as is represented in text-
figure 150, of which the four median were placed alternately
with regard to each other, and not as in earlier segments.

The gonads are visible in sections rather early in the body.
I found them without any trouble in the rather wider segments
which follow immediately upon the anterior sixty or so segments
which form the anterior section of the chain of proglottides.
A somewhat leaf-shaped mass of cells, the apex directed posteriorly,
and which represents presumably the yolk-gland as well as the

* Loe cit.

SOME MAMMALIAN TAPEWORMS. 633

ovaries, reaches from near to the anterior boundary of the pro-
glottid to beyond the middle of a segment. The éestes occupy
a restricted area in the posterior region of the segment, and
I observed in these young proglottides something like 20 or 30 of
them. They are not in contact with the ovaries, but, like them,

Text-fig. 150.

Transverse section of a posterior segment of Oochoristica sp., showing
scattered embryos embedded in general parenchyma.

e. Embryo. ¢. Longitudinal trunks of water-vessels.

are median in position. I occasionally observed also 2 or 3 testes
on one side of the ovary. Further back, when the genital ducts
first appear, the testes still form a mass which is chiefly posterior

634 MR..F, E. BEDDARD ON

to the ovary, but which has also to some extent grown round the
sides and lies laterally of the ovary.

Examining, as an entire object stained and preserved in balsam,
some segments at about the same stage of development as
those last referred to, it will be noticed that the small and
numerous testes occupy in their extension from side to side of
each proglottid about half of each proglottid. The lateral areas
not invaded by the testes are together about equal in diameter
to the middle region where the testes lie. The testes do not
reach as far as the lateral excretory vessels, and naturally, there-
fore, not to the nerve-cord.

In the mature proglottides the generative organs have been
studied by me chiefly by means of transverse sections. The
ovaries may be said to be double and le anteriorly ; they occupy
a good deal of the available space in the medullary region, and
are by no means confined to the middle of the proglottid. The
yolk-gland is single and lies behind the ovaries. The shell-gland
lies medianly in the segment on the opposite side from the ovaries.
I could find no reeeptaculum seminis, and the narrow vagina runs
a straight course for some way before its external opening.

The testes in the mature segments occupy the posterior region
of each proglottid ; but they also extend forward on either side up
to the level of the anterior border of the yolk-gland, and even
a little beyond, so that in transverse section some ovarian tissue
is occasionally seen in the same field with one or more testes. In
transverse sections, the male gonads are seen to occupy pretty
nearly the whole of the medullary layer of the worm, and are
frequently in contact with each other. Insegments of individuals
which have apparently been stretched during life the testes were
in a single row only dorsoventrally, or, at most, arranged here and
there in the form of a “ W,” one testis being slightly dorsal of
another. In such proglottides I never counted more than ten
testes in a single row. On the other hand, in proglottides that
were rather contracted than extended, the testes were frequently
for a considerable extent of the proglottid in two rows, one above
the other. In such cases as many as fourteen or even fifteen
testes could be seen in one transverse section. The testes are of
fair size, and in proglottides which are depressed occupy the
whole of the medulla from above downwards. In the other case
mentioned above, the medulla accommodated two testes one above
the other, and not much room was left unoccupied by these bodies.
In fully mature segments the testes are all of approximately the
same size, and there must be some fifty or so in a single proglottid.
Tt is easy to trace a thin membrane surrounding each testis, and
completely shutting it off from the parenchymatous tissue of the
medulla from neighbouring testes. This is the state of affairs
that we find in fully mature segments, in which, however, there is
as yet no scattering of the eggs in the parenchyma, such as occurs in
the terminal segments of the body. In a young specimen mounted
entire, the testes were quite plain in certain anterior segments of

SOME MAMMALIAN TAPEWORMS. 635

the body, wherein the genital ducts had not attained their full
development. In such a segment the testes are seen to be
distinctly more numerous than in such segments as have just
been described, a transverse section would show fully twenty
testes in a given plane (in the region of the proglottid where
they are most abundant). At the same time, these gonads
are much smaller. I have sections of proglottides which show an
intermediate state of affairs, and in which the posterior testes are
large and apparently full-er own, while anteriorly the testes are
very small and quite similar to those of immature or only in-
completely mature segments. These immature testes had no
membrane that I could detect. It appears to me that the advance
in growth of these gonads is achieved by the inclusion within a
common membrane of areas of testicular growth which have
separately originated and which in the very “youngest stages can
be seen to consist of a single cell only. It will be observed that
the disposition of the testes of this species is quite like that of
some other forms belonging to this genus as described by
Janicki *

Several fortunate sections have enabled me to see very clearly
the arrangement of the vas deferens and its mode of termination.
This tube is loosely coiled not far from its entry into the cirrus
sac. Before it enters the latter it passes in a straight course and
enters at the very extremity of the sac. It is noteworthy that
this duct is quite double the width of the vagina at its opening
into the genital cloaca and for some distance behind this point.
'The vas deferens when it enters the cirrus sac at once swells out

into a small vesicula seminalis, which thus lies within the cirrus

sac instead of outside as in most cases. The vesicula does not
by any means fill even the end of the cirrus sac in which it lies.
Immediately after this the duct narrows again, and again imme-
diately swells out into asmaller dilatation. After this comes the
narrow cirrus itself. The walls of the cirrus sae are loose and
muscular.

Liihe, the founder of this genus, in some notes upon the
anatomy of species of Oochoristica from Lizards (for example,
Tenia tuberculata of Rudolphi), remarks that the development of
the uterus must be extraordinarily rapid, as so often no inter-
mediate stages are obvious between a fully developed ovary and
the scattering of the mature eggs in the parenchyma. I quite
agree with Liihe, for the reason that the uterus is not always to
be found and is at least by no means so characteristic of this
particular Tapeworm as it is of many others. There are, how-
ever, stages to be observed. I have never seen more than a small
elongated sac lying near to either the ventral or the dorsal side of
the segment. In longitudinal sections this sac shows a tubular
form, amg 3 is, Indeed, so narrow a tube that on first observing it I
mistook it for one of the excretory vessels, and imagined that the

* Zeitschr. f. wiss. Zool. Ixxxi. 1906.
+ “ Oochoristica nov, gen. Teniadarum,” Zool, Anz. xxi. 1893, p. 650.

636 MR. F. E. BEDDARD ON

gonads were passed tothe exterior through those tubes. However,
the uterus is actually present in this form.

From the above account of the external characters and internal
anatomy of this worm we may deduce the following brief
resumé :—

Length 80-GO mm., diameter 1 mm.—2 mm. Posterior region of
the body markedly differentiated in fully adult examples by its
great thickness, but anterior end not so thin as in many forms.
Terminal segment with deep slit-like depression always present.
Necolex unarmed with rudimentary rostellum. Suckers unarmed
and sunk within the head, which is tetragonal in section. No neck,
segmentation beginning at once ; the anterior five or six proglottides
wider than scolex and seeming to form a hood into which it can be
partly retracted. Proglottides variable in proportions of length to
breadth. In many examples all of them, save the last few, broader
than long. In others some of anterior segments longer than broad.
Dorsal and ventral excretory canals not parallel, the two dorsal
being nearer together than the two ventral ; in addition to these, two
lateral narrower canals on each side. Cortical layer as thick as
medullary. Genital pores alternate regularly, the preponderance
being on one side, near to anterior border of proglottid. Testes
numerous, posterior to and at the sides of ovary. Ovaries in
anterior part of proglottid commencing shortly after anterior border.
Genital ducts pass between dorsal and ventral water tubes. Vas
deferens coiled; no obvious seminal receptacle or seminal vesicle.
Uterus sac-like, with branches ; ultimately disappearing, the eggs
being imbedded singly in the parenchyma. Occasionally traces of
uterus in form of sacs contaiung two or three ova persist.

Hab. Small intestine of Tamandua tetradactyla.

We may now consider the systematic position of this Cestoid.
The lateral position of the genital pores, the unarmed scolex, the
four suckers without appendages or hooks, the anterior position
of the cirrus pouch, the absence of more than a suggestion of a
pseudoscolex, show that this worm is to be referred to one of the
three families Anoplocephalide, Hymenolepidide, or Teeniadz,
of Ransom’s systematic table * (which is with slight differences
the same as that of Fuhrmann?’). From many of the numerous
genera contained in the first two of these families and the very
few genera of the last, the present worm is to be differentiated
by the following assemblage of characters :—scolex unarmed, no
neck; genital organs one set to each proglottid, with irregular
pores; testes numerous, posterior in position ; uterus disappears ;
eges imbedded singly in parenchyma.

The characters of the worm rather suggest the Anoplocephalide,
particularly, of course, the unarmed scolex and the absence of a
“neck.” But there is no genus in this family to which it can be
referred. The nearest is Zinstowia, in which the genital pores
are alternate, the eggs are imbedded singly, and the cortical layer

* Bull. U.S. Nat. Mus. no. 69, 1910.
+ Zool. Jahrb. extra vol. x. 1908.

SOME MAMMALIAN TAPEWORMS. 637

of the body is thick. But in Linstowia the testes extend through-
out the proglottid, the genital ducts pass ventrad of the excretory
vessels, and there are other differences. Of the Hymenolepide
the following genera only have an unarmed seolex and a single set
of generative organs with alternate pores, viz., Catenotenia, Oocho-
ristica, Rhabdometra, Anonchotenia, Metroliasthes, and Nemato-
tema. Nematotenia may be set aside as only showing strobi-
lisation posteriorly. In Anonchotenia the testes are anterior
and the eggs finally pass intoa paruterine organ. In Rhabdometra
and Metroliasthes the testes are posterior and at the sides of the
ovary, and the genital canals pass between the excretory vessels
as in the species which forms the subject of the present com-
munication ; but in those genera, as In Anonchotenia, there is a
paruterine organ into which the eggs finally pass. There remain,
therefore, by this process of exclusion only Catenotenia and
Oochoristica. 'The former genus must be eliminated, since the
genital ducts pass dorsad of both excretory tubes* and the uterus
is persistent. The present species is therefore to be referred to a
new genus or is to be included in Oochoristica. More recent
information about this latter genus than was available to Ransom
when he drew up his table shows—what is, indeed, also apparent
from that table—that the worm with which I have occupied my-
self is an Oochoristica and does not need a new genus for its
reception. Ransom’s definition is: ‘ Scolex unarmed, without
.rostellum. Single set of reproductive organs in each segment.
Genital pores irregularly alternate. Testicles numerous, sur-
rounding female glands posteriorly and on the sides. Vas
deferens coiled; seminal vesicle absent. Uterus breaks down
early and eggs become enclosed singly in egg capsules.” TI shall
now deal with the question of the species to which these worms
should be referred.

So far as I am aware, only two Tapeworms have been described
from the gut of Tamandua tetradactyla. The first of these is Tenia
tetragonocephala of Bremser, described by Diesing ft, and later, as
well as more fully, by Lihe {, whose description disagrees in
several particulars with that of Diesing. The most important
external disagreement concerns the scolex, which is represented
by Diesing as having no neck, while Liihe finds a neck 2 mm. long.
This is obviously a matter of some importance ; and I am inclined
therefore to regard the Tapeworm described here by myself as
not identical with 7’. tetragonocephala for that reason alone. With
the general anatomical description added by Lithe to Diesing’s
account the worms studied by myself fully agree, and are clearly
of the same genus which Janicki§ more recently has shown to
be Oochoristica. Janicki’s memoir contains also additional facts

* This point is not referred to by Ransom in his definition of Oochoristica ;
I have not accidentally omitted it.

+ Denkschr. k. Akad. Wien, xii. 1856.

t Arch. f. Naturg. 1895, p. 199.

§ Zeitschr. f. wiss. Zool. Ixxxi. 1906, p. 524. See also Zschokke, “ Das genus
Oochoristica,” ib. vol. \xxxiii.

Proc. Zoou, Soc.—1911, No. XLV. 45

638 MR. F. E. BEDDARD ON

upon the anatomy of O. tetragonocephala, which confirm my
opinion that the species examined by myself is not the same. In
particular, I refer tothe much more complicated excretory vessels,
which are illustrated by a text-figure *.

The second species inhabiting the gut of Tamandua tetradactyla
is Oochoristica wagenert of Janicki, who had, however, only a
single not very well-preserved example to work upon. Unfortun-
ately one very important point, as I think it, viz., the condition
of the neck, was not ascertained, and, as I understand, could not
be ascertained by Janicki from the imperfection of his specimen.
There are some other points in which my specimens were not
identical with that described by Janicki. He described ripe
proglottides as longer than unripe ones. But this is not always
the case in my specimens, since I have found quite early in the
strobila long segments. But the examination of other examples
might have led Janicki to alter this statement. For, as I have
myself shown, there is some variation in the form of the pro-
glottides in different regions of the body and in different examples.
T have noted in describing the excretory tubes certain differences
from the apparently regularly arranged six tubes found in O. wa-
genert; but here, again, it is possible that the examination of more
material would have shown that these differences do not exist.

On the other hand, I am disposed to see differences in the uterus
in the two forms. Janicki describes that organ in O. wageneri
as an irregular sac lying anteriorly in the segment and ex-
tending back as far as or beyond the ovaries, and gives a figure
showing this arrangement. The uterus in my examples was much
more irregular, and here and there, as I have described, were
quite tubular portions of it.

Even in the very last segment of the body, which might be
supposed to be fully mature, the uterus had by no means quite
disappeared in all specimens, although the majority of the eggs
were strewn through the parenchyma, as Janicki states to be the
case in his species, and as is characteristic of the genus. In other
respects I can find no differences between the facts as I read
them and Janicki’s descriptions. I do not, therefore, from a con-
sideration of all the facts, come to the conclusion that the present
species is certainly distinct from O. wageneri, but that equally
it is not certainly identical with it. Since we know that two
undoubtedly different species are found in the Edentate Tamandua,
there is no @ priori reason against the existence therein of a third
or even of more species of this genus. I prefer, therefore, in view
of these doubts, to give no name to the worm which is here
described.

Bertiella cercopitheci, sp. n.

An example of the Green Cercopitheque, Cercopithecus calli-
trichus, which died in the Society’s Gardens in February 1911,
contained two examples of a Tapeworm, which were found,

* Loc. cit. p. 535, fig. 5.

SOME MAMMALIAN TAPEWORMS. 639

contrary to what is more usual, in the colon and not attached
to the walls of that gut. It may be, therefore, that the parasites
had been loosened from their attachment in the small intestine
and had drifted into the colon, of which portion of the ali-
mentary tract they are therefore not really inhabitants. However,
both specimens were quite alive and exhibited writhing move-
ments, and it is possible therefore that they are really parasitic in
the colon. ‘They were of about the same size, and shrunk con-
siderably on preservation in alcohol. One of the two specimens,
which I have preserved entire and regard as the type of the
species, measures in the alcoholic condition 150 mm. It is re-
markable for the extreme shortness and great width of the
segments, and agrees in this particular with species of the genus
Bertiella, of which other anatomical characters prove it to be
a member.

The anterior extremity (as is shown in text-figure 151) is very
minute, and the body gradually widens up to a diameter of some
10 mm. Its general appearance is thus not precisely, although it
is generally, like that of other species of thisgenus. Accompany-
ing the increase in width of the proglottides there is also an
increase in thickness, and the posterior end of the body is about
2-3 mm. thick. In addition to the two specimens of the worm
there was a detached piece, possibly of one of these, of about an
inch in length. It appears to me that, as in Bothriocephalus, for
example, the proglottides are not shed singly but in groups, The
head of the worm is black in parts, the arrangement of the pigment
being peculiar, as I shall describe shortly. This black-headed
condition suggested to me that we might be dealing here with
examples of P. van Beneden’s Tenia melanocephala*, a parasite
from another species of African Monkey. The other characters
given by van Beneden are in perfect harmony with this view of
the identity of the species, but, as Blanchard 7 has pointed out,
the characters given are really not enough to determine the genus
to which Tenia melanocephala belongs, let alone the species.

Nor does my discovery here recorded of a black-headed Tape-
worm found in an African Monkey, and clearly referable to the
genus Fertiella, in any way settle the point at issue. For, in the
first place. I have found in a species of Dawainea (or, at any rate,
an allied genus) the same distribution of the pigment in the head
that will shortly be described in the species now under consider-
ation ; so that the mere presence of pigment in the head is clearly
no criterion of the identity of the worm. In the second place,
another species of Lertiella, viz. b. nvucronata, also from a monkey,
has been described $ in which the head is likewise pigmented. It
will be shown later that my species is not Bertiella mucronata.
Moreover, there is no reason, owing to the defective description
of van Beneden, for the identification of B. melanocephala with
B. mucronata. Thusit is necessary, as [ think, to give new name

* Mém. sur les Vers intestinaux, Paris, 1859, p. 162.

+ Mém. Soc. Zool. France, 1891, p. 186.

{ Meyner, “ Zwei neue Tzenien aus Affen,” Zeitschr. f. Naturw. 1895, p. 1.
45*

640 MR. F. E. BEDDARD ON

to the present species, in spite of the fact that it may prove
ultimately to be identical with Bertiella melanocephala.

Tn any ease, I shall be adding some further facts to the general
anatomy of the genus by the following account of this form.

Text-fig. 151.

Bertiella cercopitheci, nat. size.

A few of the posterior segments are shown more highly magnified.

The head is rather narrower than the ensuing body, in which
the proglottides are evident from the very first, there being thus no
neck. In the two specimens which I have examined, the suckers
presented different conditions. In one they were apparently

SOME MAMMALIAN TAPEWORMS. 641

absent altogether, being really retracted almost completely within
the head ; there were four little tags only protruding from the
anterior extremity. This scolex I have examined further by
transverse sections. The second worm showed only two cup-like
suckers, with their cavity directed upward as in other species of
Bertiella, One of these is distinctly larger than the other, and

Text-fig. 152.

Bertiella cercopithect.

Transverse section through scolex.

S. Sucker. P. Pigment-sheath of the same.

I presume that the remaining two were completely retracted
within the head. In a series of very nearly accurately transverse
sections through the head of the first individual, the earliest
sections showed four equidistant grooves lined by a continuation
of the thick cuticle of the scolex, which expanded towards the

642 MR. F. E, BEDDARD ON

middle of the interior of the scolex into oval chambers lined by
an equally thick cuticle. Further down the canals lost their con-
nection with the exterior and were T-shaped, with still a very
thick cuticular lining and no recognisable sucker structure. This
canal in each of the four quarters of the scolex continued for
some sections without any change, and then the suckers themselves
became plainly visible, being thus entirely retracted within the
scolex and removed from the exterior. The cavity of the suckers
is here triangular, with the sides closely pressed together. The
outlines of each sucker ave sharply marked off from the sur-
rounding tissue of the scolex by a layer of black pigment granules,
which pigment is also found in the central portion of the scolex.
The lining cuticle of the suckers is much thinner than that which
covers the scolex, and lines the canal of invagination leading to
the suckers. The outline of the scolex is here approximately
circular.

The generative organs are visible in an immature condition
very early in the body, though I have not made an accurate deter-
mination of the exact segment in which they first occur. At first
I could find no trace of any testes, simply a mass of generative
blastema which occupies the position of the future ovaries,
vitelline gland, and shell-gland, from which leads a solid rod hardly
narrower towards but not to the edge of the proglottid ; this latter
is, of course, the vagina, &e. This mass of tissue lies just anterior
to the transverse vessel, uniting the two ventral excretory vessels,
and is therefore some way from the posterior boundary of the
proglottid. In transverse sections through these very anterior
and immature proglottides it is seen to lie upon the ventral side of
the transverse vessels and to cross it obliquely to the dorsal side,
whence it passes towards the edge of the proglottid to the dorsal
side of both dorsal excretory tube and nerve. This is the same
on both sides of the body, the generative organs being single and
alternate in this worm.

Only a segment or two further back than those just described
the testes become visible, though, of course, at first quite immature.
They form a row generally only one deep (when viewed in longi-
tudinal horizontal section) extending from the excretory tubes of
one side of the body to those of the other. I counted about fifty
small testes in such a row. Here and there the row is two deep.
I could detect no trace of the vas deferens. The row of testes was
anterior to the rudimentary female organs. These latter are by
this time somewhat more developed. They still present, however,
a perfectly straight line, but reach very nearly to the edge of the
proglottid. There is, however, no external aperture. The fact
that the vagina is a tubular formation is beginning to be evident,
and the shell-gland, with radiating cells, in which it ends is plain ;
the ovary and vitelline gland lie below it and thus not in the
same plane with the vagina.

The female organs extend over more than a quarter, but not
quite a third, of the diameter of the proglottid. In still later
proglottides the uterus is for the first time quite visible and can

SOME MAMMALIAN TAPEWORMS. 643

Text-fig. 153.

Bertiella cercopitheci.

Sections through sperm-duct and vagina at three points near to their external
orifices, to illustrate relative position and structure.

C. Cirrus. Sp. Sperm-duct. Va. Vagina.

644 MR. F. E. BEDDARD ON

be traced for a considerable distance either way as a solid cord of
cells of a fibrous appearance, being more darkly stained than the
surrounding parenchyma. I could detect nolumen in this young
uterus. It runs straight along the ventral side of the row of
testes; arrived near to the lateral vessels, it bends at right angles

Text-fig. 154.

Transverse section through immature segment of Bertiella cercopitheci.

D. Dorsal water-vessel. O. Ovary. TT. Testes. U. Uterus.
V. Ventral water-vessel.

and runs straight dorsally to the inside of, and close to, the water-
vessels, crossing the vagina at right angles on its way. I could
not ascertain definitely the opening of the sperm-duct into the
cirrus sac which lay parallel with and dorsally to the vagina. Nor
could I see any branches running from the sperm-duct to the

SOME MAMMALIAN TAPEWORMS. 645

individual testes at any point. It is for this reason that I regard
the cord of cells as the beginning of the uterus and not asa sperm-
duct, which it undoubtedly suggests in its appearance and
position.

In mature segments, when both the vas deferens and the re-
ceptaculum seminis contain sperm, but when the uterus is still of
moderate dimensions only, the ¢estes are seen to extend right
across the body, when there is room for them, up to the lateral
excretory vessels on either side. In transverse sections they are
seen to be not more than two or possibly three deep in the middle
region of the proglottides. They are rather more dorsal than
ventral in position and are above the uterus. In the lateral
regions of the segments they are more crowded and often rather
closely pressed together. I could not find that the testes existed
outside of the medullary region of the segments, a point which
J mention particularly, since Meyner* met with testes (in
B. conferta) which had traversed gaps in the transverse mus-
culature and had taken up their position among the longitudinal
muscles.

The ovaries in the young proglottides are quite distinct and even
some way apart from each other, being joined by a thin bridge
which is the oviduct and which dilates in the middle into a round
sac marking the point of junction of the two oviducts. Even in
young stages the ovaries are divided into numerous digitiform
processes, which are not in the young stages thicker at their free
extremities. These processes radiate out from a common centre,
fanning out away from each ovary respectively. The ovaries are
ventral in position and radiate out towards the dorsal side of
the segment. In mature proglottides the ovaries are apparently
nearer together, that is, the connecting bridge is thicker and forms
a continuous junction of germinal tissue between them. The
processes of the ovary are now club-shaped, 7. e. thicker at their
free ends. Their arrangement is otherwise the same, but they
are much more numerous. In such proglottides the two ovaries
together occupy about one-quarter of the breadth of the body.
They are massed towards the pore side of the proglottid. The
double character thus remains distinct and is more marked than
is figured by Meyner for his species Bertiella mucronata and
B. conferta.

The vitelline glands also exhibit a double character and are to
some extent a copy of the ovaries in their general form and rela-
tions to each other. Each gland in the mature proglottides lies
a little above and between the ovaries. The dorsal position of the
vitelline glands with regard to the ovaries is shown by the fact
that the latter, when fully developed, extend dorsally at their
lateral margins and thus come to encircle the vitelline glands
lying between them. Hach vitelline gland is lobate, with numerous
oval lobes tending towards a club shape radiating out from a

* ‘Toc. cit. p. 93.

646 MR. F. E. BEDDARD ON

common centre, there being thus a likeness to the ovaries. The two
glands are likewise united by a bridge of tissue. Both the ovaries
and the vitelline glands are in close contact with the walls of the
receptaculum seminis. The shell-gland lies again rather dorsally
to the vitelline glands; but here the growth of those glands
laterally causes the shell-gland to lie rather between than above
the vitelline glands. It is in close relation with the median end
of the receptaculum seminis, which does not extend beyond it.

The sperm-duct in the mature proglottides has a form apparently
like that of other species of Bertiella parasitic in Monkeys, and is
not to be distinguished, so far as I can see, from that of Bertiella
mucronata. The sperm-duct is of considerable width from the very
first, 7. e. where it emerges from the cirrus sac. It is probable, how-
ever, that this region is really to be looked upon as representing
the vesicula seminalis of other Cestodes. A part of this dilated
sperm-duct lies actually within the cirrus sac as is depicted for
Bertiella polyorchis by v. Linstow. This portion, which is quite
short, is wider than the section which follows. The latter, however,
is also wide and lies in pretty regular coils of three alongside the
vagina as far as the commencement of the wide receptaculum
seminis. It is gorged with sperm and its walls are thin, but very
plainly recognisable by their dark staining. The coiling com-
mences directly after the emergence of the sperm-duct from the
cirrus sac, and we have therefore here a coiled region of the
sperm-duct which corresponds to that characteristic of many other
Tapeworms, but with the addition that it is the vesicula seminalis
part of the sperm-duct which is coiled.

The sperm-duct appears to come to abrupt conclusion at about
the commencement of the wide receptaculum seminis. But in
favourable sections it may be traced further as a very slender
tube closely adherent to the ventral wall of the receptaculum.
In the posterior segments of the body, which are distended
with ova, the sperm-ducts do not degenerate; on the contrary,
they are somewhat larger than in the first mature segment. They
contain, moreover, more sperm, which has somewhat distended
them: the walls thus appear thinner. The ducts in this region
of the body lie quite as coiled as in the more anterior proglottides ;
it might be supposed that they would be straightened out by the
tension caused by the enclosed spermatozoa. Nor has the sperm-
duct in any way shifted from the normal position, lying, as it does,
alongside of, and in close contact with, the vagina, which has under-
gone in this region of the body considerably greater alterations.
The same triple arrangement of the coils is visible—that is to say,
in a given transverse section there are usually three tubes to be
seen, this being, of course, the expression of the coiling.

The vagina of this species is specialised into several regions, as
it is shown to be in B. mucronata and B. conferta in the figures
of Meyner. The proglottides, from an examination of which I
have compiled the following description, appear to be in much
the same stage of sexual development as those figured by

SOME MAMMALIAN TAPEWORMS. 647

Dr. Meyner *, for which reason, of course, I have selected them.
The terminal region nearest the external pore has a thick muscular
sheath and the lumen is narrow. This section widens abruptly

Text-fig. 155.

Transverse section through mature segment of Bertiella cercopitheci.

n. Nerve-cord (only one of the three strands visible). O. Ovary. RS. Receptaculum
seminis. WV. Ventral water-vessel. Va. Vagina, to the left of which is seen
the dilated and coiled sperm-duct.

* Loe. cit. pl. i. fig. 3, pl. il. fig. 9.

648 MR. F. E. BEDDARD ON

into a thin-walled and much wider section, the muscular walls of
which are not more than, if so much as, half of the diameter of
those of the previous section. The transition is not regular, as
is figured by Meyner in the two species examined by him. The
diminution in thickness of the muscular layer, however, is
rather gradual at the orifice of the terminal part of the vagina
into the middle region.

This middle region of the vagina contained sperm in the
mature segments. It apparently ends abruptly on the median
side, but is really connected by a very narrow tube, not wider than
the sperm-duct, with the distal section of the vagina, which may
be termed the receptaculum seminis. This section is twice the
width of the last, and into it opens the duct from the ovaries &c.,
which is about as wide as the intermediate passage connecting
the receptaculum and the vagina. The receptaculum seminis
thus begins and ends abruptly. It contains sperm, as does the
distal section of the vagina. At the orifice into the genital cloaca
the sperm-duct and the vagina lie obliquely with reference to each
other. The sperm-duct is anterior, but also dorsal to the vagina,
and a little further back is completely dorsal to it. In the very
posterior segments, which are otherwise filled with the distended
uterus, the vagina with its receptaculum seminis shows an alter-
ation. It has increased in size, owing to its being gorged with
sperm.

I presume, at any rate, that the granular, in places fibrous-
looking, contents of the vagina are sperm, though they have a
different appearance from, and stain differently to, the obvious
spermatozoa which fill the neighbouring vesicula seminalis. In
the posterior proglottides the vagina shows no differentiation into
regions such as have just been described, excepting the proximal
muscular region near to the external pore. The rest formsa wide
uniform tube, and in longitudinal horizontal sections is seen to be
as wide as the proglottid is long or nearly so. In exceptional
cases even this amount of differentiation in the vagina is lost and
the wide tube pushes itself as far as the external pore, crushing
the cirrus sac into a mere rudiment.

From the above detailed description of this Tapeworm there
may be abstracted the following réswmé of its characters :—

Length of about 150 mm., greatest breadth 10 mm. Shape an
elongated cone, gradually diminishing to scolex. No neck, the
strobilisation commencing immediately after scolex. Proglottides very
short and wide, not increasing posteriorly in length. Scolex with
four suckers looking wpwards, completely retractile into head, with
black pigment. No armed rostellum. Dorsal and ventral excretory
tubes at first subequal, later the ventral very much the larger ; the
two tubes are superposed dorso-ventrally, with a transverse trunk
connecting the two ventrals. Genital pores alternate, frequently
with regularity. Generative ducts dorsal to water-vessels and nerve.
Testes numerous, forming a layer two or three thick, reaching com-
pletely between water-vessels, anterior and dorsal in position.
Cirrus feeble, no sperm-sac (vesicula seminalis). Sperm-ducts

SOME MAMMALIAN TAPEWORMS. 649

open to side and in front of vagina and pass back along ventral
margin of rows of testes. In fully mature segments sperm-ducts
increase in volume. Ovaries double, ventral and posterior in posi-
tion, in front of transverse excretory vessel, formed of nwmerous
club-shaped masses radiating from common centre. Vagina
unusually wide, showing a marked receptaculum seminis in less
fully mature proglottides ; the width increases enormously in the
fully mature proglottides until it is nearly as wide as the proglottid
is long. Genital cloaca short. Uterus single, sac-like, without
definite outgrowths, but somewhat irregular in form, filling most of
the proglottid. The eggs without V-shaped apparatus, with a
thinner inner shell and a very wide and lax outer membrane.
Proglottides apparently not detached singly, but in groups.

Hab. Cercopithecus callitrichus.

It is clear from this definition that the Tapeworm now under
consideration has been rightly referred by me to the genus
Bertiella. There is, I think, no other genus which shows the
same assemblage of characters as those which I have just set forth
in brief. It remains to be seen what position the species occupies
within the genus, of which we now know a good many different
species.

It seems that we may select the following characters as
distinctive of this species, which will be thus definable :—

BERTIELLA CERCOPITHECI, Sp. n.

Length 150 mm. or more, breadth 10 mm. posteriorly, gradually
tapering to head. Scolew with black pigment scattered throughout
middle of scolexw and forming a special layer round each sucker.
Suckers directed anteriorly, and completely retractile. No neck,
strobilisation commencing at once. Lateral nerve-cord divided into
three, the middle one the largest. Lateral water-vessels superposed,
the dorsal much the smaller, Testes extending between water-vessels,
forming a layer two or three deep and three or four wide in middle
region of proylottid, more numerous laterally. WVesicula seminalis
very long and coiled, extending into cirrus sac. Vagina divisible into
two well-marked regions and communicating by very narrow interval
with wide receptaculum semis. Hggs without pyriform apparatus.

Hab. Cercopithecus callitrichus, in colon (2).

Following the subdivision of the species of Bertiella by
Zschokke * into three groups, the present species evidently
belongs to the first group, to which Zschokke assigned Tania
mucronata and 7. conferta of Meynery. Bertia studeri and
B. satyri of Blanchard} are doubtfully admitted by Braun § into
the genus Bertiella as recognised by the better-known species fully
described by Meyner, for Blanchard does not describe the genital

* “Neue Studien an Cestoden aplacentaler Satigethiere,” Zeitschr. wiss. Zool.
Ixv. 1897, p. 404.

+ “Zwei neue Tenien aus Affen,” Zeitschr. f. Naturw. 1895, p. 1.

¢ Mém. Soc. Zool. France, 1891, p. 186.

§ Bronn’s Klassen u. Ordn. des Thierreich, iv. Abth. 18, p. 1712.

650 MR. F, E. BEDDARD ON

organs, except in so far as to remark upon their alternating pores
and upon the structure of the ova. Nor does he say anything of
the pigmentation of the head, which is so marked a feature of my
species and of the otherwise unrecognisable Tania melanocephala
of van Beneden *.

B. mucronata was described from an American Ape, JJycetes
niger, and I gather from Meyner’s description that the Tapeworms
themselves were brought from Paraguay, and thus real inhabitants
of the Monkey in question, which, therefore, had not been
infected in a menagerie. This species shows the same pigmenta-
tion of the head as mine, and its general shape? is not very
different. Nevertheless, it appears to me that in the species
described in the present paper the anterior end is more tapering,
and has a very small diameter for a longer stretch. Indeed, the
general outline of the worm described in the present paper is
more like Meyner’s B. confertat, which is, moreover, a parasite
of an Old World Monkey, Macacus radiatus, and therefore,
perhaps, more likely on @ priori grounds to be identical with mine.
I think, however, that B. conferta may be put out of court in this
comparison, for the reason that no mention is made of a black
pigmentation in the scolex, which is so clear a feature of my
species and of B. mucronata. Nevertheless, 5. conferta agrees
with my species and differs from 4. mucronata in that the vagina
enters the receptaculum suddenly, there being no gradual widening
as in B. mucronata, where the tubes are continuous. The suckers
are said, however, to be lateral in position, as in B. mucronata §.
This is one of the principal reasons which lead me to regard my
species as distinct. Of B. mucronata Meyner writes :—“ Sind
nicht vollstiindig nach vorn, sondern mehr zur Seite gerechtet.”
In a series of transverse sections through B. cercopitheci, which I
have described above, it is very clear that the apertures of the
suckers are not lateral nor slit along the retracted sucker, such as
is so obvious in the other species described in the present paper.

Nor does Meyner describe the remarkable pigment-sheath to
each sucker which characterises my species. Furthermore, it
appears that B. mucronata has a “neck”; for Meyner writes ||:—
“ Schon etwa 0:9 mm. vom Skolexscheitel entfernt, also in einem
Stadium, wo der Kérper noch keine Spur einer Gliederung
erkennen lisst, gruppiren sich,” etc. Stiles, in reviewing 4 the
characteristics of this species, points out that B. mueronata is
also to be defined by the fact that the generative canal passes to
the exterior between the nerve-cord on the one hand and the
two excretory vessels on the other. My sections of B. cercopitheci
show very plainly that the generative canal passes dorsally to
both nerve and lateral excretory vessels. This is particularly

* Mém. sur les Vers intestinaux, Paris, 1859.

+ Loe. cit. pl. 1. fig. 1. t Loe. cit. pl. ii. fig. 8.

§ In other Bertiella (e.g. in Sluiter’s B. plastica, see Centralbl. Bakt. xix. 1896,
p- 941) the suckers look upwards.

|| Loe, cit. p. 81. € Proc. U.S. Nat. Mus. 1896, p. 145.

SOME MAMMALIAN TAPEWORMS. 651

obvious in young proglottides, which are so slender that the
disposition of these structures can be seen in a single section.

A final point of systematic importance concerns the horn-like
processes of the innermost egg-shell, which are figured by Meyner
and stated by him to be always very obvious. I have been abso-
lutely unable to detect these structures, even when using an oil
immersion-lens (4 in. Leitz). We must, therefore, agree that
this group of species of Bertiella is to be characterised, as the
two remaining groups into which Zschokke divides the genus, by
‘“birnformiger Apparat nicht constant.” Iam unable to compare
the species which I describe here as Bertiella cercopitheci with two
species deseribed by Gottheil * from Macacus cynomolgus and from
that species and Cebus capucinus, since they are not sufficiently
diagnosed, and Stiles is of opinion that they are only doubtfully to
be referred to the genus Dertiella. Indeed, the position of the
genital pores is not referred to.

Another Monkey parasite with which my Tapeworm might be
compared is v. Linstow’s Bertiella polyorchis + from Macacus cyno-
molgus. This is interesting, from the point of view of the ege-
shells, and confirms what I have said above concerning the absence
of the horn-like processes in 4. cercopitheci ; for it is hardly likely
that so experienced a helminthologist as Dr. von Linstow would
have overlooked these structures were they present, and his figure
of the egg of 4. polyorchis does not show them. Furthermore,
the suckers look forward and a dilation of the sperm-duct within
the cirrus sac is figured. The species, however, differs, as I
believe, from Lertiella cercopitheci by its less complicated vagina,
by the much greater number of testes which fill the middle of the
proglottid, and by the absence of any black pigmentation in the
head. At any rate, the latter point is not referred to.

Thysanosoma gambianum, sp. n.

I obtained from an example of the Gambian Pouched Rat,
Cricetomys gambianus, which died in June 1909, a considerable
number of Tapeworms which I regard as being of a new species
and belonging most probably to the family Anoplocephalide, The
Rodent had been one year and ten months in the Society’s
Gardens before its death, and it is, therefore, quite credible that
it was infected with these Cestodes when it arrived in London.
The material, as well as being abundant, was well preserved, and I
am therefore able to give a fairly comprehensive account of the
anatomy of the species, which presents certain new combinations
of characters.

The external characters alone appear to place this worm in
either the genus Anoplocephala or Zschokkeella. There are no
other genera in which the scolex is unarmed, the genital pores
are unilateral and the neck is absent, and the segments until the

* Journ. Comp. Med. & Surgery, 1887, vol. vii. The species are not named and

are referred to Tenia.
+ Arch. f. Naturg. xxi. 1905, p. 270,

652 MR. F. E. BEDDARD ON

very end of the body are broader than long. Unilateral genital
pores occur only in these two genera among the subfamily or
family Anoplocephalide or Anoplocephaline. There are, however,
several genera among the remaining Tetracotylea which possess
genital pores in sequence upon one side only. But the number
of genera is small and other external characters prevent a con-
fusion. Thus the Tapeworm which forms the subject of the
present section cannot be confused with Aploparaksis, Diorchis,
Gryporhynchus, Paruterina, Culcitella, Oligorchis, Lateriporus,
Dilepis, Davainea, Progynia, Idiogenes, Chapmannia, Pro-
orchida, Monopylidiwm, or Cyclorchida, since all of these have
a circle or more than one circle of hooks upon the rostellum.
There remains Hymenolepis, which is to be distinguished by

Text-fig. 156.

Thysanosoma gambianum, nat. size.

To the right are a few segments more highly magnified to show genital papilla.

possessing a “neck,” which the present species does not, and by
the limited series of testes in each proglottid. I am thus correct,
as I believe, in regarding this worm as a member of the Anoplo-
cephalide. ‘

There are reasons both for and against referring the Tape-
worm from the Gambian Pouched Rat to either of these genera
or to a new genus, into which I shall enter at length after
detailing its anatomy. This species isa large worm, measuring

SOME MAMMALIAN TAPEWORMS. 653

some 6 inches in length. A very marked characteristic is the
fact that the segments are wider than long throughout almost the
whole of the body. It is only at the extreme end that they
become longer. The segments overlap successively, so that it is
possible to detect the anterior margin of each segment. The
worm is not very stout, but flattened after the typical Tapeworm
fashion ; the most posterior segments alone being rather thicker
in a dorso-ventral direction. The diameter of the body in the
middle is some 6 mm. There is no marked colour, though, as will
be seen presently, there is some. internal pigmentation. The
general character of the segmentation suggests that the pro-
glottides are not shed.

This species like Anoplocephala has no “ neck,” as it is termed,
that is to say, there is no undifferentiated zone following the scolex,

Text-fig. 157.

Scolex of Thysanosoma gambianum.

where the limits of the proglottides are unrecognisable. The
definite proglottides are obvious from the very first. They are
at first rather narrower than they become posteriorly; but the
diameter of the body very soon attains to its full dimensions.

Proc, Zoou, Soc.—1911, No. XLVI. 46

654 MR. F, E, BEDDARD ON

The cirrus of each segment is not visible, or at least not con-
spicuous, when the worm is viewed with a lens, in a protruded
condition. This is due to the small size of these organs, which
will be more fully described later. I have not found it possible
to differentiate the ventral from the dorsal surface. The scolex, as
already stated, is unarmed. I investigated this part of the worm
by the section method, and am therefore able to state positively
that there is no rostellum nor any trace of hooks. Nor do the
suckers show any hooks that I could detect. I am convinced
that they are absent altogether from the scolex of this worm. A
regular series of transverse sections across the “ head ” shows a
rostellar region above the region of the suckers, which is oblong
in section ; it contains no pit or depression of any kind that was
obvious to me. The suckers are the usual four and as usual
symmetrical. When the scolex is viewed in its entirety under a
low power of the microscope the suckers seem to face rather
upwards ; but they do not lie on the upper side of the head as in
some Tetracotylea. There is only a slight obliquity. The orifice
of the sucker in such a preparation is very plain and rounded.
There is no appearance of a slit-like orifice such as is described
in the present paper in Oochoristica, and certain details in the
minute structure of the suckers in these two genera show corre-
sponding differences. I find in two series of transverse
sections of the suckers of the two Tapeworms, that in the present
species eleven or twelve sections displayed the entire sucker,
of which only five showed its cup-like orifice; in Oochoristica,
on the other hand, the entire sucker required seventeen sections
of equal thickness for its display, of which twelve or thirteen
showed the obviously more slit-like orifice.

As in Oochoristica the orifice of the external integument is
much more limited than that of the sucker itself. The two coin-
cided for only the space of two sections, rather less than in
Oochoristica. ‘This, however, I take to be simply a measure of the
state of contraction of the suckers. It is also important to notice
that the outer integument is grooved for a little space anteriorly
to the appearance of the sucker itself, which suggests a con-
firmation of a view already put forward, and that is the inde-
pendence of the suckers of the more specialised Tapeworms from
the bothria of the Dibothriata. It appears to be possible that
this grooving is the persistent trace of the bothrium, upon which
the sucker has been added as a subsequent development.

In series of transverse sections through the head there is
another feature of the suckers which deserves mention. It is
very clear from such sections that the growth of the sucker
has taken place in a definite direction.

Although I am unable to give histological details, it is certainly
the fact that anteriorly the sucker is in organic connection with
the general tissues of the head. ‘There is no break; it begins
gradually with a recognisable condensation of tissue marked by
its deeper staining with carmine. On the other hand, at the

SOME MAMMALIAN TAPEWORMS, 655

posterior end of the sucker its tissues come to an abrupt end, and
I invariably found a shallow cavity between the sucker and the
medullary tissue of the head. This suggests that the original
position of the sucker is more apical and that it is directed
upwards—a, state of affairs which is actually seen to persist in
many species of Tetracotylea.

The orifices of the generative organs are, as already stated,
completely unilateral. I have examined pieces of the worm
mounted whole, and many series of sections, and find that in
every case there is a regular sequence of the apertures which lie
upon one side of the body*. These orifices are not conspicuous
on examination with a lens, as they are in many Tapeworms, and
for two reasons. In the first place, the orifices are actually small,
and in the second place, I have never observed thecirrus to protrude.
Thus it is necessary to make a microscopical examination in order
to detect the genital pores. Apart from microscopic sections,
where, of course, it is easy to find the pores, I have only observed
them (that is, in pieces of the worm mounted entire) in the wider
posterior segments. Here they are seen to lie at just about the
middle of the proglottides. The orifices project slightly and are
quite circular.

The excretory system of this Tapeworm presents certain unusual
features. In transverse sections through some of the middle
segments of the body there are four longitudinal canals perfectly
obvious in the body and no more than four, unlike, for instance,
the genus Oochoristica, with which the present genus has some
characters in common. These four tubes are lateral in position,
and are all of them at about the same level. It is thus impossible
by position to distinguish the dorsal from the ventral excretory
canals. The two on each side are at a considerable distance apart.
The distance which separates the outer from the inner of the two
is about as great as that which separates the outer tube from the
nearest margin of the body. The tubes can, however, be dif-
ferentiated by their size. The outermost of the two excretory
canals is not more than one-third of the diameter of the inner-
most tube. The form is not, however, so small that it can be
missed in transverse sections. In the very anterior segments I
have noticed a third lateral vessel on each side; and in this region
of the body, moreover, the two main tubes have a greater incli-
nation respectively to the dorsal or ventral side. There is an
abundant plexus of vessels connected with these and traversing
the proglottides, but 1 am unable to give details. I think,
however, that I have seen numerous external pores.

The testes have a somewhat peculiar arrangement, which is
distinctive of this worm. The bulk of these very numerous
gonads lie upon that side of the body upon which the generative

* T cannot determine whether this is right or left with certainty, since in trans-
verse sections the gonads are not definitely dorsal or ventral in position, 7. e. nearer
to one surface of the proglottid or the other, while the dorsal and ventral excretory
tubes are parallel to each other and lie in the same plane with the nerve-cord.

46*

656 MR. F. E. BEDDARD ON

pores are not situated. They are naturally in the medullary part
of the body, and in transverse sections are seen to occupy the
greater part of this, being neither dorsal nor ventral in position,
but simply central. They reach towards the margin of the
segment which is nearest to them, far beyond the outermost of
the two excretory tubes of their side of the body—ain fact, up to
the nerve-cord ; medianly they do not reach the median line of
the body, but extend beyond the innermost of the two excretory
tubes. They occupy perhaps, when seen in this view, one-fourth
of the entire breadth of a proglottid. They are not very close
together and are, at most, in two rows dorso-ventrally, there being
thirteen or fourteen to each row, and thus something like thirty
may be visible in a single section. In longitudinal sections seven
or eight of these double or partly single rows are seen, and they
may be observed to occupy the greater part of the segment.
There is also a second set of testes at the opposite margin of the
proglottid, very much fewer in number—perhaps four or five in a
transverse row, which lie on either side of the outer excretory
tube.

The ovary may be regarded as a paired structure, and the two
are partly separated by the yolk-gland, which lies between and
behind them. In transverse sections through the ovaries, before
the other parts of the female reproductive system have been
reached, each ovary is seen to be distinct from its fellow and to
lie on either side and below the larger of the two excretory tubes,
which, in such a section, is seen to be not median in position.
The excretory vessel is here only just below the cortical layer of
the proglottid, though actually in the medullary layer. The two
ovaries are not quite in contact below the vessel, or, at any rate,
they can be recognised as two distinct bodies of a bushy form not
unlike that which has been figured in other Tapeworms. Within
a few sections from that which has served as the basis of the
above description the shell-gland is seen taking the place of part
of that ovary which lies median of, 2.e. not to the pore side of,
the excretory vessel, and the oviduct leading from it to the ovary
of that side is conspicuous. In a section or so further on in the
series the yolk-gland appears; although, as already said, this
gland divides the two ovaries, it does not lie symmetrically with
reference to the excretory tube. It lies almost entirely on the
median side of this tube but also below it, though it does not
extend at all on to the pore side of the excretory vessel. It
occupies nearly the whole of the space on the inner median side
of the excretory tube that, in previous sections, is occupied by the
ovary of that side. In the next proglottid to that whose ovaries
and associated glands have just been described, the ovary to the
outside reached nearer to the outermost and smaller of the two
laterally placed excretory tubes, but without arriving at it. In
this case it is important to notice that there was no accessory
group of testes lying between the ovary and the outermost of
the two excretory vessels such as occur in the proglottid first

SOME MAMMALIAN TAPEWORMS. 657

described, and which are referred to elsewhere in the description
of this species. Series of longitudinal (sagittal and horizontal)
sections brought out the position of the ovaries and their relation-
ship to adjacent glands still more clearly. Such sections also
showed that the gonads are quite in the middle of the proglottid,
2.e. dorso-ventrally. It should be remarked that the double
character of the ovary was not always so strongly marked, and
that it sometimes lay entirely upon the pore side of the more
median (and larger) excretory vessel. Furthermore, I have also
observed the yolk-gland to lie upon the median side of the larger
excretory tube as well as upon the pore side. There is thus some
variability.

It is, perhaps, noteworthy that the gonads of the pore side,
which consist of the ovary, yolk-gland, and the testes, together
occupy about the same space as the gonad and testes only, in this
case, of the opposite side of the proglottid. The female gonads, at
any rate, are visible very early in the body, within twenty seg-
ments, I dare say, of the head. Shortly after their commencement
the mass of tissue which is to form ovary, yolk-gland, and, as I
presume, oviduct, vagina, &e., is seen very plainly to extend to-
wards the periphery of the proglottid between the two longitudinal
excretory vessels, a feature of systematic importance in this group
which it is necessary to note. In the sexually mature proglottides
the oviduct runs straight from the ovary as a thin-walled delicate
tube not easy to see, which is sometimes wider, at least near to
its termination, in the ejaculatory apparatus, runs to near the
distal termination, and then suddenly narrows into a very fine
tube with thick, darkly-stained walls which, after a very short
course, again suddenly widens into a long sac which is as wide as
the cirrus sac and runs beside it and below in position. This
terminal reservoir is laxly surrounded by what appears to be an
adventitious sheath of muscular fibres, which have a circular
direction and thus appear cut across; they are very obvious
through their deep staining. This layer of muscles was some-
times, but not always, observed to commence with the very thin
region of the vagina. The wide terminal region of the vagina
bends towards the cirrus sac and again becomes narrow, opening
in common with the latter into the genital cloaca, which is very
short. This genital cloaca is quite distinct from an ingrowth of
the outer layer of the body which meets it and forms the actual
pore; this has been described in considering the external
characters. The cirrus sac, as already implied, is neither wide nor
long. The cirrus was never seen in a protruded condition, but
always lay a darkly staining rod within the pouch. The vas
deferens, directly it leaves the cirrus sac, is thrown into a large
and complicated coil, which reaches for a considerable distance
into the interior of the body. I think that it ends in a dilated
vesicula seminalis above the ovary; there is, in any case, a dilated
pouch in this region full of sperm, which does not appear to have
anything to do with the female ducts.

658 MR. F. E. BEDDARD ON

Tn sections through the posterior segment of the body, the ripe
eges, with their sacs, are seen to occupy the whole of the available
space, that being, of course, the medullary region of the proglottid.
The segments are, in fact, stuffed full of eggs, and, in accordance
with this, their dorso-ventral diameter has somewhat increased,
though not to so very great an extent as in some other Tapeworms,
for instance, in certain specimens of the Oochoristica described in
the present paper. A closer examination of the eggs shows that
they are imbedded a few together in a dense and darkly staining

Text-fig. 158.

Transverse section through proglottid of Thysanosoma gambianum to show
numerous paruterine organs (e). ¢. Water-vessels.

mass of tissue, which closely invests them. These sacs appear to
me to be, without doubt, the equivalents of the paruterine organs
of many Cestoidea. They are not precisely sacs, in that there is
no central lumen occupied by the eggs; they are rather concen-
trations of the medullary tissue round a series of eggs. These
bodies are of approximately equal size and contain much the same
number of eggs or, rather, embryos. There is no question here
of a circle of paruterine organs surrounding a centrally placed

SOME MAMMALIAN 'TAPEWORMS. 659

uterus. The uterus has entirely vanished, and the paruterine
organs are imbedded in the tissues of the body.

Text-fig. 159.

Paruterine organs of Thysanosoma gambianum more highly magnified.

S. Walls of paruterine sacs. O. Embryos.

So much for the condition in the fully ripe proglottides. Harlier
in the body this formation of paruterine sacs is preceded by a
uterus which is not very conspicuous, and consists of not much
more than a transversely running tube extending nearly right
across the proglottid in which the eggs occur, but with which the
uterus never appears to be stuffed. I could find no outgrowths
of this centrally placed uterus, and there was certainly nothing
in the nature of a reticular formation of its cavity. Gradually
the cavity of the uterus appeared, as it were, to dry up and the
eggs were found—to continue the simile—stranded in the tissue of
the body. At this time the formation of the paruterine sacs
became visible. Round each egg, or round two or three, as the case
might be, the tissue of the medullary region of the proglottid
became denser, this being shown by its darker staining. There
was thus a concentration of tissue round the ova. This concen-
tration of tissue had no relations that I could detect with the

660 ON SOME MAMMALIAN TAPEWORMS.

uterus. There was nothing like the formation of diverticula of
the latter walled by the condensed parenchyma.

The characters of this worm may be briefly summed up as
follows :—

Scolex unarmed, with no rostellum, only a raised area. Strobila
commencing without an intermediate neck. Proglottides much
wider than long, except at the very end of the body. Proglottides
very numerous, the length of worm being some 6 inches with a
greatest diameter of 6 mm. Genital pores unilateral. Eacretory
tubes four, parallel to each other. Cortical layer of body as thick
as medullary. Testes chiefly massed upon the side of the body
Surthest from the genital pore, very numerous, median in position.
In addition, a small number of similar testes on either side of the
outermost excretory vessel of the pore side of the proglottid. Vas
deferens with a large coil and a vesicula seminalis above ovary ;
cirrus sac not long. Genital cloaca small, with circular muscles.
Ovaries double, on either side of innermost of excretory tubes of pore
side of segment. Yolk-gland on one side of same excretory vessel
behind ovary. Shell-gland nearer to the middle of the body above
the yolk-gland. Seminal receptacle, long and not much swollen, |
begins soon after the terminal chamber of the vagina. Uterus broad
and sac-like, occupying a great deal of the middle of the proglottid.
Many paruterine organs present in later stages.

The characters given in the above paragraph are not distinctive
of any known genus of Tetracotylea. And I am, indeed, disposed
to think that ultimately it will be necessary to form a separate
genus for this worm from the Gambian Pouched Rat. In the
meantime, however, I do not burden zoological nomenclature with
an additional name until the possibility of its distinctness becomes
more settled. Besides it is also possible that the existence of this
species removes a barrier between the two genera Thysanosoma
and Anoplocephala. Until I had become aware of the numerous
paruterine organs, I was disposed to refer the worm to Anoplo-
cephala, with which genus it obviously has many points in
common. But the existence of these characteristic paruterine
organs—and in such great numbers—is a reason for removing it
from Anoplocephala and uniting it with Thysanosoma. On the
other hand, the latter genus has either double or single sets of
generative organs, and, correspondingly, either two pores upon each
proglottid or alternating pores, while the Tetracotylean described
in the foregoing pages has generative pores all upon one side.
Nevertheless, the double set of testes seems to be a last trace of an
originally completely double set of gonads and ducts, such as occurs
in some proglottides of other Thysanosomas. If the small set of
the testes existing in this species upon the pore side of the ovaries
were to disappear it would be, as I think, impossible to separate
this genus from Anoplocephala or Zschokkeella ; but the definition
of those genera would have to be enlarged in order to take in the
numerous paruterine organs, which is, after all, perhaps the chief
reason for referring this worm to the subfamily Thysanosominz
which is mainly thus characterised.

ON THE WHALEBONE WHALES. 661

30. On the Natural History of Whalebone Whales.
By J. A. Morcu, (Christiania) *.

[Received March 3, 1911: Read April 4, 1911.]
(Text-figures 160-163.)

No period in the annals of modern whaling exhibits such an
intense activity as that which has been developed during the
course of the last two years, after the pioneer expeditions of the
preceding years to antarctic, subantarctic, and other waters
in the Southern Hemisphere had demonstrated the wealth of
Cetacea which is to be found in some of these localities. The
rise in the price of oil to figures which, through shortage in the
crop of oil-seeds, have not been obtained for the last twenty-five
years, has had a further stimulating effect upon the development
of this pursuit in those southern waters.

The situation to-day, then, is that, after the short run of six
years, whaling in the Southern Hemisphere has attained a com-
mercial importance entirely overshadowing that of the industry
in our northern latitudes, which is now more than forty years old.
As an example, it may be mentioned that last season, from
the island of South Georgia alone, fourteen whaling steamers
brought 106,800 barrels of oil, which is more than the world’s
total production of whale-oil three years ago !

Kven during the latter part of the last century the attention
of Norwegian whalers was directed to these distant regions; and
in 1893 two expeditions started from Norway to try their luck
in the chase of Seals and Right Whales in southern seas. One,
in the ‘ Antarctic,’ proceeded to Australian waters, and the other,
in the ‘Jason, to the islands south-east of Cape Horn. The
pecuniary results were not encouraging, and further attempts
were given up for the time. Then came the Nordenskidld scientific
expedition in 1901, and Capt. C. A. Larsen, in the ‘ Antarctic’
belonging to that expedition, had an opportunity of continuing
his observations made during his previous voyage in the ‘ Jason.’
Having satisfied himself of the seemingly limitless numbers of
whales in those waters, he succeeded, in 1904, in interesting parties
in Argentina in his plan for establishing a whaling-station on the
island of South Georgia; and he began operations there about
the new year, 1905.

Meanwhile, in 1903, the Norwegian Storthing had passed a
law prohibiting whaling on the northern coasts of Norway, and
the now homeless whalers had to go in search of other fields for
their operations. In the spring of that year a comparatively small
vessel was fitted out as a floating factory and dispatched to
Spitzbergen as an experiment. This proved successful; and in
the next summer a larger vessel proceeded to these waters,

* Communicated by S. F. Harmer, Sc.D., F.R.S., V.P.Z.S.

662 MR. J. A. MORCH ON

with the same result. The floating factory, which was going to
play such an important part in the development of modern
whaling, became hereby an established fact; and in October 1905
this same vessel, together with two whaling steamers, was dis-
patched to the Falkland Islands and South Shetland. Upon the
vessel’s successful return with a nearly full cargo of oil in June
1906 the ice was broken ; and now followed with intense activity
the fitting out of a number of floating factories, as also of build-
ings and appliances for the erection of land stations in various
localities in the Southern Hemisphere.

Text-fig. 160.

~

Floating factory and whaling steamer in harbour, Deception Island,
South Shetland ; with floating carcases of Humpbacks.

From the southern coasts of South America, South Shetland,
the Falkland Islands, South Georgia, the coasts of South Africa,
and Kerguelen Island, whaling is now being prosecuted with an
ever-increasing number of whaling steamers and with returns of
oil undreamt of only a few years ago.

The species which are principally hunted are the Humpback
Whale (Megaptera boops), the Blue Whale (Balenoptera sibbaldi),
and the Finback Whale (B. musculus) ; and, in one locality (the
Falkland Islands), the “Seihval” or Rudolphi’s Whale (B. borealis),
which also occurs on the coast of Chili and the west coast of
South Africa without having been actually hunted there yet.

THE WHALEBONE WHALES. 663

The Sperm Whale (Physeter macrocephalus) and the Southern
Right Whale (Balena australis) are occasional visitors in some of
the localities; the latter species having been observed nearly
every year trekking in schools along South Georgia and towards
the Patagonian coast. At South Shetland, Bottlenose Whales
(Hyperoodon rostratus) have also been observed in small schools,
but have not been the objects of pursuit. The appearance of
this species near the coast in Bransfield Strait is interesting,
but as none were caught no observations could be made upon
what constitutes their food in this locality.

Humpbacks constitute the great bulk of the Whales caught in
most of the above-mentioned localities. The cow of this species
is supposed to go pregnant from ten to eleven months, and,
judging from their numbers there, must be the most prolific of
Whales. In February 1910 I observed at South Shetland two
Humpback feetuses about 23 feet long which, if we accept the
supposition of the late Prof. G. Guldberg as to the growth of
Humpback fcetuses, should be about 21 months old (impregnation,
therefore, about the first of November). At South Georgia,
L am informed, even at the commencement of the season, Hump-
backs with calves from about 12 feet long are met with, and
foetuses are also found here in their first months of development.
According to these observations, the indications are that the
Humpback Whales in these southern latitudes may be supposed
to give birth to their young at some time in September or
October, as against April and the neighbouring weeks in our
northern latitudes.

I may add that from the circumstance that in some of these
southern localities the Whales are only flensed and the carcases
left to be driven by wind and tide, it is unfortunately only by
the merest chance that observations upon the period of gestation
of the various species can be made. As, however, a more appro-
priate disposition of the huge masses of meat and bones may
be looked for in the future, opportunities should also present
themselves for acquiring enough material for examination in
this respect.

An interesting phenomenon is observed from South Georgia
relating to the Humpback Whales. At certain times all the
Humpbacks that are brought in have the belly nearly white ; this
variety may then disappear and those caught for some time may
have the belly marbled ; schools with their bellies entirely dark
may then put in an appearance, succeeded by the first variety, and
so on. At South Shetland I also observed these several varieties
of the Humpback, although they did not there appear in distinct
schools, but mingled. The bulls were here in a decided majority,
and the individuals observed in February and March were mostly
young.

A locality which is attracting serious attention at present 1S
the western coast of South Africa. During the months May to
October especially, the Humpback Whales have been observed in

664 MR. J. A. MORCH ON

great numbers and mostly off the coast of Portuguese West
Africa. I think the question may reasonably be raised whether
we have not here the great bulk of the Humpbacks from the
South Georgia region on their annual migratory route? In this

Text-fig. 161.

Humpback Whale (nostrils open).

locality, which is interesting in more than one respect, observa-
tions might probably be made upon fcetuses of Humpback Whales
in the later stages of their development. Large schools of
Rudolphi’s Whales and Blue Whales have also been observed

THE WHALEBONE WHALES. 665

along these coasts, feeding upon plankton, but the few Blue
Whales caught were all very lean.

The coast of Chili, from which whaling is at present being
prosecuted, also offers opportunities for interesting observations
upon the periods of gestation of various species of Whales, especi-
ally those of the Blue Whales and Rudolphi’s Whales, which are
very little known.

From a consular report which has come to my notice, it appears
that great numbers of the Finback Whale congregate in the
waters along the Brazil coast between South latitudes 12 and 18
every year during the period from May to November. Until
more definite information is obtained upon this subject, further
comment must, however, be reserved.

In the Northern Hemisphere, modern whaling is at present
being prosecuted from the following localities :—Spitzbergen,
Iceland, the Faeroe Islands, Shetland, the Hebrides, the western
coast of Ireland, Newfoundland (one station also in Labrador),
British Columbia, Japan, and Korea, while preparations are also
being made for an early start in Alaska and in the Sea of Okhotsk
from Saghalien Island.

When the veteran whaler Svend Foyn had perfected his
harpoon-gun in the latter part of the sixties, and had commenced
operations in Varangerfiord on the Finmark coast, the Blue
Whales were his only objects of pursuit; and, so long as the
hunting was carried on on a moderate scale, continued to be so for
a series of years. As, however, after the expiration of his patent
an increasing number of new companies entered the field, the
other species of Whales—Finbacks, Humpbacks, and occasional
schools of Rudolphi’s Whales—also became the objects of the chase.
Successively, however, the Blue Whales became scarcer and scarcer,
and the other species of Whales, especially the Finbacks, came
to play the principal part in the catches. The same order of
things has, as a rule, repeated itself also in most other whaling-
grounds in the North Atlantic.

As observations upon the breeding-season and the period of
gestation of the various species of Whales in our northern latitudes
can only be made during a limited part of the year, no exact
information may be said to exist bearing fully upon these ques-
tions. The results from the Finmark coast may very briefly be
summed up thus :—

The Blue Whales have often been observed in coition during
the summer ; the indications are that this act may take place also
at other times of the year. They have never been observed with
small calves in that locality, but occasionally with larger ones.
The period of gestation is supposed to last more than twelve
months, and it is believed that there is no fixed time of the year
when they give birth to their young; this probably takes place
in American temperate waters*. Contents of the stomach of

* The Sulphur-bottom variety has occasionally been obseryed in Finmark waters.

666 MR. J. A. MORCH ON

various Blue Whales have been observed by Prof. G. O. Sars to
consist chiefly of Boreophausia inermis. The Blue Whales are
remarkably free from parasites, this perhaps owing to the cir-
cumstance that their outer skin easily peels off.

Pinback Whales.—Impregnation is supposed to take place in
January—March, and the period of gestation to be about twelve
months. The length of the new-born calf is about 20 feet. Very
young calves are always seen together with full-grown individuals
in schools, older calves sometimes in schools by themselves. It is
supposed in the locality in question that among 50-60 Finback
cows 10-15 will have feetuses and a lesser number will be
accompanied by calves. Some cows give birth to their young
in those waters, some go westwards to other localities. It is
supposed that the cows do not become pregnant every year.

Text-fig. 162.

Typical site of Epizoic Crustacea on the ventral surface of a Humpback Whale.

As the Finback Whales in the North Atlantic feed on
plankton, caplin (J/allotus villosus),and herrings, their distribution
at most times of the year depends on the appearance of these
various sorts of food, The whalers distinguish three varieties:

THE WHALEBONE WHALES. 667

the blackish, the grey, and the yellowish. The blackish forms
follow the schools of caplin and herrings. The time of ap-
pearance of the two other varieties seems to indicate that they
feed chiefly on plankton. Further information upon the
migratory route of these varieties would be interesting, and
observations on the subject might be made during the progress
of the herring fisheries. I may add that at South Shetland, in
February and March, 1910, I observed several Finback and Blue
Whales which were covered by a muddy, yellowish deposit
which could easily be scraped off. As I had not’a microscope
with me, no information upon the nature of this substance
could be obtained. Penmnella is, so far as I know, the only
parasite that has been found, and this only occasionally.

Text-fig. 163.

Epizoic Crustacea (Coronula diadema and Conchoderma auritum)
from Humpback Whale.

Humpback Whales —During the early months of the year in-
dividuals of this species are met with on the Finmark coast going
west. They are at that time followed by large calves, are restless,
and approach the shores. About April the cows are supposed to
give birth to their young in subtropical waters. Impregnation
is supposed to take place shortly afterwards. The Humpbacks
arrive again in the waters north of Finmark in summer;
and around Bear Island they are found in July feeding on
plankton (Boreophausia inermis, &c.) and caplin, and are then
seen with calves about 20 feet long. The fcetuses found at
this time of the year are about 20 inches long. In September
they go east into the Barents Sea, but observations upon their
life during the last months of the year are wanting. They are

668 MR. J. A. MORCH ON

supposed then to feed on caplin and to follow the schools of these
fishes near the “ East Ice” and move westwards with them during
the winter. Those killed on the westward trek have been found
to have empty stomachs.

On account of the tough outer skins of the Humpback, parasites
can easily fasten themselves thereto, and this species is, as a matter
of fact, especially infested by various forms :—Coronula diadema,
Conchoderma auritum, and Paracyamus boopis. Pennella is rarely
found. ;

Rudolphi’s Whales (B. borealis) are very erratic in their
appearances. In 1884, for instance, only six were killed on the
Finmark coast; in 1885, 659. This is a typical plankton Whale,
and it appears on that coast only during the summer, feeding on
Calanus &c. The foetuses have a length of from 3 to 4 feet in
June, from which it may be inferred that the cows give birth to
their young during the latter months of the year in localities
at present unknown.

In Shetland in 1906 I observed on a Rudolphi’s Whale, which
had the front end of its lower jaw deformed, a colony of Concho-
derma auritum fastened thereto. ‘This is the only instance of
parasites on this species that I know of.

Plankton being the only or principal subsistence for the Whales
in question, an exceedingly interesting problem is suggested :—
What part do the great ocean currents play as highways and
feeding grounds for these Whales during their annual migratory
route ?

From investigations carried on by Prof. Nansen and _ his
assistants it has been proved that the cold polar water has a
beneficial effect upon the vegetable life in the open sea. The
cold polar currents, by mixing with waters of a higher temperature,
create favourable conditions for the growth of plankton and
higher marine life.

From these investigations, then, we may infer that it is along
the border layers of the great polar currents where these meet
and intermingle with warmer currents or waters that, given a
sufficient actinity of light for the production of vegetable plankton,
we may expect to find the most favourable conditions for the
subsistence of the plankton Whales.

We know from our northern latitudes that the waters
along the northern coast of Iceland, Finmark, and along the
western coast of Spitzbergen have attracted and been able to
maintain for a long series of years a considerable or even a very
great number of Whales.

On the contrary, we have seen that in localities which are under
the principal influence of a cold polar current—for instance, the
coast of Newfoundland—the stock of Whales has in the course
of only a very limited number of years been seriously reduced,
although the number of whaling steamers employed would not
have been excessive had the same favourable conditions in the sea
prevailed as, for instance, along the northern coast of Iceland.

THE WHALEBONE WHALES. 669

The considerable number of Whales which were met with along
the coast of Newfoundland during the earlier years of hunting
may probably have accumulated, slowly, during the course of time
by natural multiplication, z.e. the older ones have been followed
by their young and these also have come back to the same
locality. If the catches, then, had been proportional to the
approximate natural increase, the industry might most probably
have been carried on to the same extent for a long time to come.

It may be taken for granted that the great bulk of the Whales
which during the spring months migrate northwards through the
Atlantic take an easterly direction and Spread in a fan-like
manner towards Iceland, the Faeroe Islands, Shetland, Spitzbergen,
and the northern coasts of Norway. Now the question is: Does
the western part of the Atlantic bordering upon Nova Scotia,
Cape Breton, and Newfoundland offer them any inducement in the
form of sufficient food which they may be supposed to find at this
time of the year growing along the border of the Gulf-Stream in
various localities? This may, I think, be answered in the negative ;
and I believe that we have here also circumstances which may
have contributed to the condition of things which has manifested
itself off Newfoundland.

On the Murman Coast, also, there has been snails instance
of a similar case. In the eighties, two whaling stations were
started here by the Russians, and under apparently favourable
auspices as to the number of Whales which were seen in this
locality. After a few seasons of successful hunting, however, the
Whales became scarcer and scarcer, so that at last their pursuit
had to be given up. This agrees also very well with our own
experience on the Finmark coast, where the whaling stations in
the eastern localities had to be moved westwards at a com-
paratively early date, as the Whales became scarcer in the colder
areas towards the Murman Coast.

If the statistics of the Whale fisheries in some of our northern
localities are examined, it will strike an observer that the years of
good or poor returns generally run into periods of years of either
the one or the other kind. There may one year be foggy or
boisterous weather accounting for smaller catches, but these
obstructions do not generally last in periods of years. It will take
a closer study of the hydrobiological conditions of larger areas
during the particular years to give a satisfactory explanation of
this phenomenon, and I am only here indicating its existence.

With the latter part of August, or the first part of September,
most of the Whales in our northern latitudes have left their
summer haunts. The migratory routes of the Blue Whales and
the Rudolphi’s Whales between the seasons, and to some extent
also during these, may well be said to be shrouded in mystery.
The two American bomb-lances, which in the years 1888 and
1898 were found in Blue Whales on the coast of Finmark, do not
give us much clue as to the particular locality where they had
been fired into the animals.

Proc, Zoot, Soc.—1911, No. XLVII, AT

670 ON THE WHALEBONE WHALES.

Of the Finback Whales a considerable number must stay in our
home waters also between successive summer seasons, following
and feeding upon the shoals of caplin and herrings at various
localities and times of the year. But their numbers are not so
ereat as to account for the large schools which migrate to southern
latitudes. In the Mediterranean considerable numbers of Fin-
back Whales are seen, but further information from this locality
is lacking.

Tn addition to what has been mentioned above with regard to
the Humpback Whales, nothing is known about the migratory
route of those which visit the waters off Newfoundland and
northwards. It would be of interest to know if they follow
the caplin in these localities, like the FEST p/AGES in the Barents
Sea.

In the month of May, Humpbacks are found off the Azores,
Bermuda Islands, and occasionally the Antilles. In 1899 parts
of an American bomb-lance were found in a Humpback on the
Finmark coast.

The part played by the Gulf-Stream in the biology of Whales
in the North Atlantic is paralleled by the influence exerted by the
Antarctic current or great West Wind Drift in the South Sea.
The localities which have shown themselves to be rich whaling
grounds, such as the coast of Chili, South Georgia, and the couiheet
coast of West Africa, are under the influence of this current.
By the intermingling ‘of the cold waters from this current with
waters from the adjacent warmer currents, conditions must be
created favourable for the production of the masses of plankton
which are sometimes seen in these localities. What part the
Humboldt current along the western coast of South America and
the Benguela current along the western coast of South Africa
play as migratory routes and as feeding grounds during several
months of the year for those species of Whales which at the
present time are of the greatest commercial importance among
the Cetacea, must be left to future investigations to disclose.

The geographical positions of the various whaling grounds in
the Southern Hemisphere should offer special opportunities for
observations upon the migrations, breeding-season, food, and other
questions of biological and economical interest relating to these

Whales.

ON THE NEST OF A GREY STRUTHIDEA. 671

EXHIBITIONS AND NOTICES.
April 25, 1911.

Dr. S. F. Harmer, M.A., F.R.S., Vice-President,
in the Chair.

THE Secretary read the following report on the additions to
the Society’s Menagerie during the month of March 1911 :—

The number of registered additions to the Society’s Menagerie
during the month of March last was 270. Of these 73 were
acquired by presentation, 137 by purchase, 45 were received on
deposit, 8 in exchange, and 7 were born in the Gardens.

The number of departures during the same period, by deaths
and removals, was 158.

Amongst the additions special attention may be directed to :—

2 Siamang Gibbons (Symphalangus syndactylus) 9 2, from
Sumatra, purchased on March 10th.

2 Neumann’s Vervet Cercopitheques (Cercopithecus centralis),
new to the Collection, from Lake Kivu, Central Africa, deposited
on March 23rd.

2 Ruffed Lemurs (Lemur varius) $ 2, from Madagascar, pre-
sented by Frederick Burgoyne, Esq., F.Z.8., on March 15th.

2 Pumas (felis concolor), from Pernambuco, presented by John
Sparks, Esq., F.Z.S., on March 11th.

1 Elephant-Seal (J/acrorhinus crozetensis), new to the Col-
lection, from the Crozet Islands, presented to The King’s African
Collection by Dr. Louis Péringuey, F.Z.8., and deposited by
H.M. Tue Kine on March 23rd.

1 Gundlach’s Troupial (Quiscalus gundlachi), new to the
Collection, from the Greater Antilles, presented by the Countess
of Suffolk on March 2nd.

Mr. C. Tate Reean, M.A., F.Z.S., exhibited a series of lantern-
slides of scales of the Salmon (Salmo salar), and showed how the
life-history of the fish could be read from its scales.

Mr. D. Seru-Smitu, F.Z.8., the Society’s Curator of Birds,
exhibited :-—

(1) A nest (text-fig. 164, p. 672) of the Grey Struthidea or
Apostle Bird (Struthidea cinerea), composed entirely of mud, and
built on a branch in the Western Aviary.

(2) Lantern-slides from photographs of the King Penguin
(Aptenodytes pennanti) and Black-footed Penguins (Spheniscus
demersus) Showing the method of moulting.

(3) Lantern-slides of a number of wild Swainson’s Lorikeets
(Trichoglossus novee-hollandie), from photographs kindly sent
by Mrs, Innes, of Mackay, North Queensland. These birds came

47*

672 ON A COLLECTION OF SKINS FROM UGANDA.

in large numbers to feed daily at a table, on syrup provided,
settling without fear upon the head, shoulders, and arms of the

lady who fed them.
Text-fig. 164.

Nest of the Grey Struthidea.

Some Mammals from Uganda.

Dr. C. Curisty, F.Z.8., exhibited specimens from a collection
of skins of antelopes, hyrax, monkey, cheetah, serval and serva-
line cat, &e., obtained in the Chagwe forests in the south-east of
the Uganda Protectorate, and made the following remarks :—

“Perhaps the most interesting specimens in the collection
before you are those of Cephalophus weynsi. This red duiker
was first described by Mr. O. Thomas, F.R.S., from two skins
from eastern Congo. Subsequently a specimen was obtained by
Mr. L. M. Seth-Smith in the Budongo forest on the east of Lake
Albert. An imperfect skin in the National Collection, labelled
Cephalophus johnstoni, also obtained from the Lake Albert region,
and described some time ago by Mr. Thomas, may possibly also
belong to this species.

‘My own four specimens were shot in the Mabira forests more

ON A LOIN-CLOTH OF A NATIVE OF NORTHERN NIGERIA. 673

than 100 miles east of the Budonga, and with very little forest
intervening. The locality is a new one, and is probably the
easternmost limit of the species, which, after another gap of
150 miles of more or less open country still further east, gives
place to Cephalophus ignifer, the common red duiker of the East
African upland and rift-valley forests.

“The two species are closely allied. In both the body-hair is
short and close-laid, and the horns slope backward in a line with
the forehead. Amongst other characteristics C. weynsi may be
distinguished by the hairs on the nape being reversed forward.

“These sleek, heavy, short-legged duikers are very pig-like in
gait and appearance, carrying the head low. They are found only
in dense forest, and, so far as my experience goes, never even feed
in the open, unlike the grey duiker so commonly to be met with
at forest-edges and often seen in the open. My two best heads of
C. weynsi measure 44 and 43 inches in length respectively.

‘“‘T may also draw your attention to the specimens of hyrax—
Procavia emini and P. dorsalis (2). Naturalists do not seem to
quite realise that certain species of hyrax, the Dendrohyrax group,
are entirely arboreal, never living amongst rocks or holes in the
ground, but inhabiting the largest trees in dense tropical forests ;
whilst other species are rock dassies, and though able to run up
and down the face of a perpendicular rock or even to play about
the neighbouring bushes, yet are in no sense arboreal. The
members of the Dendrohyrax group do not even live in hollow
trees as a rule, but upon the branches.

‘On looking closely at the skins of these two animals, P. emane
and P. dorsalis (2), both killed in high trees, it is interesting to
observe that the long bristles amongst the fur, so numerous and
so conspicuous, especially on the hinder part of the body, in those
species which are not arboreal, are here obviously absent or
only to be found on the neck or shoulders. I find, as the result
of an examination of the skins in the British Museum, that this
distinctive peculiarity holds good for the two groups in almost
every instance, the arboreal Dendrohyrax group being almost
without them, while in the rock-inhabiting species they are very
conspicuously developed, mainly posteriorly.

“Tt, seems probable that these long stiff hairs are tactile organs
and of very considerable use in dark burrows and holes amongst
the rocks; whilst it is easy to see that they are of less utility
on the branches of trees, and in time, no doubt, would become
rudimentary or disappear altogether.

“he weird, nocturnal, ventriloquistic cries of both groups of
these animals are even more extraordinary than their powers of
climbing.”

Dr. Currsty also exhibited a loin-cloth taken in 1898 from the
dead body of a native in the Gando-Bornu district of Northern
Nigeria. In referring to it, he said :—‘‘ When first I caught
sight of this ornamental piece of wearing apparel it seemed so
peculiar that I stopped to secure it under considerable difficulties.

674 DR. WILLIAM NICOLL ON A UNIQUE

It is apparently made from the skin of a young Cobus cob; but
the white of the belly part and the inner part of the hind legs
you will see are transversely striped with inch-wide stripes, some
two or three inches apart, of a reddish-brown colouring exactly
resembling that of the animal. These stripes are doubtless made
with some native pigment or dye; but they are so placed and the
colour is so permanent, so fixed and difficult to remove by any
means, that serious doubts have been entertained, as Mr. Thomas
will tell you, as to whether they were artificial or natural
markings. one

“Tt will be also noted that the white hair on the striped belly-
parts is curled and not unlike sheep’s wool, but quite unlike any
known antelope. If this curling of the hair in this situation is
not natural, I can only suggest that it is produced by the heat
and chafing of the wearer’s thighs.”

Mr. Oldfield Thomas, in commenting upon Dr. Christy’s exhibit,
stated that there was little doubt that the stripes and curly
appearance of the hair were artificial, but the fact that the dye,
whatever it was, so closely resembled the natural colour of the
animal and appeared to be so “ fixed,” the possibility of the skin
belonging to some antelope hitherto quite unknown had been very
carefully considered at the Natural History Museum.

On a unique Pathological Condition in a Hare.
(Text-figure 165.)

Dr. Wiuit1am Nicout, M.A., F.Z.8., exhibited some pre-
parations from a Common Hare (Lepus europeus), which showed
an interesting and unique pathological condition. The hare was
obtained by W. Raphael Muckley, Esq., and sent by him to the
British Museum, whence it was forwarded to the Lister Institute
of Preventive Medicine. The manner in which the hare died
was somewhat remarkable. It was observed by Mr. Muckley to
pitch violently out of a hedge on to the roadside where it lay
struggling, and it died about three hours later. To him the
symptoms seemed to point to poisoning. At the post mortem
examination the liver was found to be extensively invaded with
small whitish chalky deposits of various sizes, especially con-
spicuous on the posterior surface. There was also a considerable
amount of chronic inflammation around the liver, with adhesions
to the diaphragm. On section. the liver was friable and gritty.
The concretions were amorphous and insoluble in acid. The bile-
ducts showed some fibrous thickening. On microscopic exam-
ination of a scraping from the liver, numerous ova of peculiar
structure were seen. They were fairly uniform in size, measuring
057 x ‘033 mm. At first sight they bore a considerable resem-
blance to the eggs of Trichuris (Trichocephalus) or Trichosoma,
but on more careful inspection they presented one or two unusual
features. The colour was much lighter, being greyish instead of
brown. The shell consisted of two layers, the outer of which
was marked with pronounced radial striations; the imner was

PATHOLOGICAL CONDITION IN A HARE. 675

constituted of fine concentrically-arranged lamelle. The two
layers were not in close apposition but were separated by a
narrow, somewhat irregular space. The egg is thus provided
with a double shell. At each pole of the egg there was a small
circular aperture, piercing both layers of the shell. The external
surface of the shell presented a well-marked papillated appear-
ance, the papille being irregularly rounded. Inside the shell

Text-fig. 165.

M.R., del.

Eggs of a Nematode worm from the liver of a Hare.

A. External surface of egg. bB. Optical section of egg: two-celled stage.
C. Egg in four-celled stage with thickened inner wall: from vagina.

there was a thin, continuous membrane investing the egg. The
polar apertures were each closed by a plug of homogeneous
material which was separate and distinct from the membrane
investing the egg-cells. They completely filled the apertures but
did not bulge beyond them. The eggs were mostly in the two-
celled stage, but a number of them showed four cells.

676 ON A UNIQUE PATHOLOGICAL CONDITION IN A HARE.

From the appearance of the eggs it is obvious that the lesions
were due to a Nematode worm of the 77ichosoma group. Search
was accordingly made and a number of worms were obtained. In
every case, however, they were fragments lacking the anterior
and posterior extremities. The longest specimen measured
27 mm. It was a female, full of eggs, and it was very slender.
It was narrowest at the anterior end (‘14 mm.), the breadth
increasing gradually towards the posterior end where it was
‘23mm. The cuticle was marked throughout with fine annular
striations. The vagina was of considerable length but no genital
aperture was seen, so that it must have been far forward. In
the vagina a number of eggs were present, which were much
larger than those first seen. They measured -070—078 x -:040—
045 mm. The shell, too, was much thicker, the increase being
almost entirely due to a great thickening of the inner layer. |
The polar apertures were somewhat reduced in size and the eggs
were for the most part in the four-celled stage.

The lesions in the liver showed much resemblance to those
seen in advanced stages of coccidiosis, a very common condition
in rabbits. For this reason they might readily be diagnosed as
such on cursory examination. This, so far as can be gathered, is
the first record of such an infection of the liver of hares or rabbits
by parasitic nematodes of this kind.

An analogous condition has not infrequently been met with in
rats, and has been reported from Europe, India, and Australia.
No record has been made of its occurrence in this country.

Whether this parasite in the hare is the same as that in the
rat must remain doubtful, for the descriptions of the latter have
hitherto not paid much attention to the character of the eggs
beyond mentioning that they resembled those of Z’richosoma.

An attempt is being made to hatch the eggs and to produce
infection in rabbits and rats, but the development is extremely
slow even at a constant temperature of 26° C.

In addition to the lesions in the liver the hare had a very
heavy infection of Z'richostrongylus retorteformis in the intestine
and a slight intestinal infection with Coccidiwm cuniculi. The
intestine and the appendix, moreover, showed a large number of
small caleareous patches, but on examination nothing of a parasitic
nature could be detected in the patches. This was of interest
from the fact that I had already seen similar patches in the
cecum of a Variable Hare, sent me from the Society’s Gardens.
The liver, unfortunately, was not submitted for examination.
That these patches might have something to do with the liver
condition was not impossible, for they might be considered as
degeneration following injury such as the passage of a worm
through the intestinal wall.

The hare, further, showed signs of recent parturition and there
was a septic condition of the uterus, and it must remain an open
question as to whether death was due to this or to the liver
disease.

Cae

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PZ.S.19U.PLXXVI.

London Stereoscopic Co. imp.

M. Rhodes, del.

LECHRIORCHIS VALIDUS.

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PZ.S.19M. PL XXVoL.

M. Rhodes, del. London Stereoscopic Co imp.

6. 7. OCHETOSOMA FORMOSUM.
8-10. DASYMETRA CONFERTA.

ON NEW TREMATODES FROM REPTILES. 677

PAPERS.

31. On Three New Trematodes from Reptiles.
By Witiiam Nicout, M.A., D.Sc., M.B., F.Z.S.*

[Received December 29, 1910: Read April 25, 1911.]
(Plates XX VII. & XXVIII.T)

The following notes were made on a collection handed over to
me by the Prosector of the Zoological Society. In no case was
the habitat noted, but from their affinities it may be presumed
that the specimens all came from the lungs, the mouth, or the
cesophagus. The specimens are interesting as forming an im-
portant addition to our knowledge of the large variety of forms
which inhabit the air-passages and anterior end of the alimentary
canal of reptiles and batrachians. To the taxonomy of this
particular group Odhner has recently (1910) made a valuable
contribution, which will be further referred to later.

The first species is from the Hog-nosed Snake (Heterodon
platyrhinus), and I include it provisionally in the genus Lechri-
orchis Stafford, 1904. Like most of Stafford’s genera, this genus
is insufficiently defined, although the fact that he includes in it
the well-described species LZ. (Renifer) elongatus Pratt, 1903, is
something to go upon. It is, therefore, necessary to amend
Stafford’s definition somewhat, as follows.

Genus LecHRiIorcHIs Stafford, 1904.

Resembling Renifer Pratt, 1903, except in the following
particulars. The intestinal diverticula extend a short distance
beyond the testes; the genital aperture is further from the edge
of the body, midway between it and the pharynx ; there is a well-
developed vagina.

These constitute, as far as appears, the only differences between
the two genera.

LECHRIORCHIS VALIDUS, sp. n. (Plate X XVII. figs. 1-5.)

This is a species of moderate size, measuring 3°6-7:-4 mm. in
length by ‘9-1-4 mm. in breadth. The smallest specimens had
just begun to produce ova, so that the minimum adult size is
probably about 3 mm. The body is elongated, the length being
4_5 times the breadth. The latter is comparatively uniform, but
the tail is distinctly pointed. The anterior end is more rounded,
while there is a tendency fora slight narrowing to occur about
or behind the ventral sucker. In young specimens the body is
flattened, but it becomes much thicker as the uterus increases in
size. In transverse section an adult specimen presents a strongly
convex dorsal surface and a fiat or slightly convex ventral surface.

* From the Lister Institute of Preventive Medicine, London.
+ For explanation of the Plates see p. 686.

678 DR. WILLIAM NICOLL ON

The cuticle is fairly thick, somewhat deciduous, and is studded
throughout its whole extent by stout salient spines. These are
regularly arranged, and become sparse towards the posterior end.
The underlying musculature has the usual formation.

In an average adult specimen of 6°5 mm. length the oral
sucker has a diameter of -45 mm. It is subterminal, globular,
and its aperture has a marked muscular rim. The thickness of
its wall is about ‘13 mm. In the same specimen the ventral
sucker measures 66 mm. It is also globular, slightly flattened,
and its wall is -18 mm. thick. The sucker ratio is, therefore,
very approximately 2:3. The diameter of the sucker relative to
the body-length diminishes as the animal increases in size.
Thus, in the smaller specimens the ventral sucker is + of the
body-length, while in the largest it is only j. The ventral
sucker is situated about 4 of the body-length from the anterior
end. In this respect, again, the young specimens differ from the
older ones in that they have the ventral sucker relatively further
back.

There is a very short prepharynx; usually the pharynx 1s
contiguous with the oral sucker, and it measures -23 x17 mm.
The cesophagus is about ? of the length of the pharynx, although
it is longer in young specimens. The intestinal bifurcation takes
place well in front of the ventral sucker. The intestinal diverticula
run almost parallel to the edges of the body. They bend in a
little just behind the ventral sucker, but they are pressed out
again by the testes. At their termination, just behind the
posterior testis, they usually turn in a little. Their ends are
about 2 of the body-length from the posterior end. They are
comparatively narrow tubes, and their wall is crmkled on the
inner side. The outer side is plain. They are lined by a layer
of low epithelium. The cesophagus has the usual cuticular lining.
Throughout their whole extent the diverticula are somewhat
ventral in position.

The excretory system is characteristic and agrees with Odhner’s
supposition as to its form in the group to which this species belongs.
The vesicle consists of a fairly broad main stem, opening at the
tip of the tail and passing forwards to the shell-gland, where it
divides into two limbs. These diverge to form a Y, and each passes
over the edge of the ventral sucker and terminates halfway
between the sucker and the intestinal bifurcation. The main stem
is pressed close to the dorsal surface, but the limbs are not quite
so dorsal in position. They eventually come to lie close up to the
intestinal diverticula and separate these from the uterus and
cirrus-pouch respectively (Pl. XX VII. fig. 4). The main stem is
much compressed dorso-ventrally, but the limbs are round or com~
pressed transversely. In addition to this central system, however,
numerous lateral twigs are given off from the stem and limbs.
These all radiate outwards towards the edges of the body, dividing
and subdividing and eventually forming an intricate anastomosis
(Pl. XX VII. fig. 3). The entire lateral fields of the body are thus
filled with a complex network of excretory tubes, and the amount

NEW TREMATODES FROM REPTILES. 679

of parenchymatous tissue is very much reduced. This is more
particularly the case in the post-acetabular region. It gives rise
to a reticulated appearance, which is strikingly seen in young
specimens (Pl. XX VII. fig. 1). In older specimens it is obscured
by the growth of the uterus.

The genital glands are difficult to distinguish in the adult, but
are easily seen in younger specimens. The testes lie not very far
behind the ventral sucker, near and internal to the ends of the ~
intestinal diverticula. They are obliquely situated, the left testis
being half its diameter in advance of the right, and they are
separated from each other by the uterus. In a young specimen,
the uterus being narrow, they lie quite close together, but they
are pressed further and further apart by the growth of the uterus.
They are flat, elongated oval bodies measuring at least *8 x 4 mm.
Their outer margin is plain, but their inner margin is indented
im one or more places, so that the outline is somewhat irregular.
At first they lie almost flat in the body, but the expansion of the
uterus pushes their inner border towards the dorsal surface so
that eventually they are considerably tilted.

The genital aperture is situated on the left side midway between
the pharynx and the edge of the body. It is always on the level
of the pharynx. The cirrus-pouch is of considerable length, and
is a conspicuous object. It is elongated, somewhat slender, and
extends to the middle of the ventral sucker. Its wall is remarkable
for the great development of the longitudinal muscular fibres,
which are stout and very prominent. The circular fibres are
much smaller. Within the pouch there is a more or less highly
convoluted vesicula seminalis. Usually it is simply bent double
(Pl. XXVIT. fig. 2), but frequently it is much more twisted. It
is small compared with the size of the cirrus-pouch. It is not
much dilated, and it is connected with the pars prostatica by
a narrow duct. The pars prostatica is of relatively great length.
It is an almost straight tube extending from a little in front of
the ventral sucker to the point where the cirrus-pouch crosses the
left intestinal diverticulum. It is uniform and fairly narrow.
Surrounding it are numerous prostatic cells which fill up the
greater part of the cirrus-pouch. The ductus ejaculatorius is
short and narrow, and the exsertile, unarmed cirrus is not very
long. The genital sinus is quite small.

The ovary is situated just behind the end of the cirrus-pouch,
over the right posterior quadrant of the ventral sucker. Half of
it lies beyond the sucker, and it frequently overlaps the adjacent
intestinal diverticulum. It is analmost globular body, somewhat
flattened dorso-ventrally, and is about half the size of the testes.
Behind and internal to it lies a compact shell-gland, with a short
ootype and a Laurer’s canal, but no receptaculum seminis.
Laurer’s canal.is short and opens dorsally in the middle line,
about the level of the ovary. A small yolk reservoir lies dorsal
to the shell-gland. The yolk-glands are of limited extent. They
are entirely lateral and lie close to the outer side of the intestinal
diverticula. On each side there are about half a dozen ill-defined

680 DR. WILLIAM NICOLL ON

groups of follicles, which extend from midway between the
intestinal bifurcation and the anterior edge of the ventral sucker
to near the ends of the intestines. The initial part of the uterus
is practically empty. In section it can be made out with difticulty
as an extremely narrow tube, passing backwards from the shell-
gland. Behind the right testis it widens out to form a recep-
taculum seminis uterinum, consisting of about four or five small
dilatations. Further back a few ova appear, but the uterus
still remains somewhat narrow. When it has nearly reached the
posterior end of the body it turns abruptly on itself to form an
ascending limb. Almost immediately this begins to dilate, and
it has only proceeded a short distance before it almost completely
fills the interior of the body. It passes forward, over and between
the testes, crosses the ventral sucker, and terminates ina straight,
thick-walled vagina, which is about half the length of the cirrus-
pouch, and lies on the left side. The uterus thus consists of
a descending and an ascending limb, the former being empty for
the greater part of its length, and the latter being enormously
dilated. The increase in size, therefore, takes place, not by an
increase in convolutions, but by a great dilatation of the ascending
limb. The ova are very numerous, rounded oblong in shape, and
dark brownin colour. They have a large well-marked operculum.
Many of them are more oval than oblong, and this gives rise to
some variation in dimensions. From a large number of measure-
ments the limits were found to be -038-:045 mm. for the length,
and -018-:023 mm. for the breadth, and the average ‘040 x :021
mm. The extreme sizes observed were 045 x-018 mm. for the
most oblong ova, and -038 x :023 mm. for the most oval.

No case of amphitypy was observed in any of the two dozen
specimens forming the collection. The nearest approach was in
one specimen where the testes were practically symmetrical, the
left being a trifle behind the right, but the ovary and genital
aperture were normal. In all the other specimens the position
of these structures was exactly as I have described. The species,
however, is extremely variable in one respect, namely, the
posterior limit of the yolk-glands. Hardly two specimens agree
in this respect. In some specimens they extend a short distance
beyond the testes, in others they reach the middle of the posterior
testis, and again in others they stop short of the testes. In
addition, they are very frequently asymmetrical, extending further
back on the right than on the left, or more rarely vice versa. The
fact, however, that their anterior limit is constantly symmetrical
induced me to consider a symmetrical posterior limit as the normal.
In two specimens also, the intestinal diverticula were of unequal
length, the left diverticulum being considerably shorter than the
right, which was normal. The position of the genital aperture
varied only very slightly, and most of the apparent variations
were due to contraction. The length of the cirrus-pouch was
practically constant, although in one specimen it extended nearly
to the posterior border of the ventral sucker. The size of the ova
was constant within the limits noted, and no increase in size takes

NEW TREMATODES FROM REPTILES. 681

place as the animal grows older. The average size of the ova in
young specimens was found to be the same as that in fully grown
specimens, and I am inclined to view with some doubt Odhner’s
statement (2. p. 59) that the ova in Renifer sauromates Poir.
increase in size as the animal grows older.

From Lechriorchis elongatus Pratt, this species is distinguished
by having more unequal suckers, the ventral being decidedly
larger, the yolk-glands being more extensive, being present some
distance in front of the ventral sucker, and in having slightly
larger eggs. From Z. primus Staff.,it appears to be distinguished
by its much smaller eggs, its smaller ventral sucker, and probably
in other respects.

« The second lot of specimens consists of five from an Annulated
Snake (Leptodira annulata). They bear a close superficial resem-
blance to Lechriorchis validus, but they do not belong to the same
genus. The only genus to which they can at present be referred
is Ochetosoma Braun, 1902, but they do not entirely agree with
the definition of that genus as given by Braun. In internal
anatomy they correspond quite closely, but they are not nearly so
flattened as O. monstruosum Brn., the only species of the genus.
The only other genus to which they could be referred is Renifer
Pratt, 1903, but from that they differ radically in the configuration
of the uterus.

OCHETOSOMA FORMOSUM, sp. n. (Plate XXVIII. figs. 6 & 7.)

The body is elongated and slightly flattened. The length is
3°7-5'7 mm., and the greatest breadth, about the ventral sucker,
is 11-16 mm. The breadth is therefore rather less than 4 of
the length. The body tapers gradually towards each end.

In an average specimen (length 4°5 mm.) the oral sucker has
a diameter of -45 mm., 7. e. ;/5 of the body-length. It is globular
and subterminal. The ventral sucker is situated 1:7 mm. ‘from
the anterior end. It is slightly oval, the transverse diameter
being ‘64 mm., and the longitudinal -57 mm. The sucker ratio
is therefore approximately 3: 4.

The cuticle is extremely deciduous; in most of the specimens
it was almost entirely stripped off. Only in one was it intact, and
even then not completely so; from this specimen the presence of
minute regular spines was determined.

There is a short prepharynx followed by an almost globular
pharynx, measuring “17x ‘16 mm. The cesophagus is about the
same length as the pharynx (:2 mm.), and the intestinal bifurcation
occurs well in front of the ventral sucker. The diverticula
diverge pretty widely, and they terminate almost immediately
behind the ventral sucker, the ends being somewhat turned in.
They are narrow and irregularly dilated, but there is no crenation
on their inner wall as in Lechriorchis validus. They are lined by
low epithelium. :

The excretory system is almost identical with that in Lechriorchis
validus, The vesicle consists of a similar dorsal main stem, which

682 DR. WILLIAM NICOLL ON

divides just behind the ovary into two limbs, which pass forwards.
a short distance in front of the ventral sucker. They are thus
not so long as in the above mentioned species. The main stem,
too, appears more expanded. There is the same system of
secondary lateral branches which anastomose freely in the sides
of the body.

The genital aperture is situated almost at the extreme left
margin of the body, on the level of the posterior end of the
pharynx. The genital sinus is very small. The cirrus-pouch is
very like that of Lechriorchis validus, but it is shorter. It is
usually disposed obliquely, and it terminates a short distance in
front of the ventral sucker, from which its end is usually separated
by a coil of the uterus. In one specimen it almost reached the
sucker. It contains a small convoluted vesicula seminalis, ending
in a narrow portion which runs into the pars prostatica. The
latter is not so long as in the foregoing species, and it 1s more
expanded, especially at its posterior end. The prostatic cells are
numerous. ‘There isa short ductus ejaculatorius, and an unarmed
cirrus of moderate length. The testes are situated just behind
the middle of the body (:3 mm. behind the ventral sucker). They
are symmetrical and lateral. They lie behind the ends of the
intestines, but are separated from them by folds of the uterus.
The length of each is about °5 mm. They are fairly thick and
elongated, and their outer margin is always distinctly divided
into three large lobes, which may in addition be slightly crenated.
The inner margins are completely obscured by the overlying folds
of the uterus.

The ovary is situated over the right posterior quadrant of the
ventral sucker and projects half beyond it. It is obliquely ovoid
and measures ‘°24x°17 mm. A large shell-gland lies close to its
inner side. The yolk-glands are entirely lateral and of limited
extent. They reach from the anterior border of the ventral
sucker to about the middle of the testes. Again in this species,
however, the posterior limit is extremely variable, and may be
anywhere between the anterior and posterior borders of the testes,
but never beyond them. The anterior limit is practically con-
stant. The uterus fills almost the whole of the post-acetabular
region, but its configuration is entirely different from that in
Lechriorchis. ' Here, again, the descending limb is small and
almost empty ; reaching the posterior end of the body it turns
into the ascending limb. In this case, however, accommodation
for the enormous number of ova is obtained not by excessive
dilatation, but by numerous convolutions, the diameter of the
uterus not being very greatly increased. The convolutions have a
markedly transverse disposition, extending from side to side of
the body. In the region of the testes the convolutions are shorter
and stouter. An additional small convolution is formed in front
of the ventral sucker. The uterus terminates in a well-marked
vagina, which is about a third of the length of the cirrus-pouch.
The ova are very like those of Lechriorchis validus, but are usually
more oval. They have a large distinctly-marked operculum, and

NEW TREMATODES FROM REPTILES. 683

they vary in length from -034 mm. to ‘042 mm. by -017 mm. to
°021 mm. in breadth. The average is about -4 x°2 mm.

In this species, again, no case of amphitypy was observed, and
the only pronounced variation was in respect of the posterior
limit of the yolk-glands as described above.

The species obviously presents a close resemblance to the genus
Renifer Pratt, sens. strict. In the shortness of the intestinal
diverticula, the symmetrical situation of the testes, and the
extreme lateral position of the genital aperture, the agreement is
complete. The essential difference lies in the configuration of the
uterus. Renifer ellipticus Pratt, the type species, is unfortunately
not fully grown and the ultimate disposition is not apparent. In
R. sauromates Poirier, the uterus is of the same type as in
Lechriorchis validus, and if this be taken as characteristic of the
genus, then O. formosum must be separated from that genus.

It is evident that the three genera Renifer, Lechriorchis, and
Ochetosoma ave somewhat closely related, and they differ from all
the other members of the family Lepodermatide in the extreme
lateral and forward position of the genital aperture. They
evidently form the nucleus of a group, but the extremely profuse
variety met with in the family renders it somewhat difficult to
divide it into definite subfamilies. Provisionally, however, these
three genera may be classified under Pratt’s subfamily Reniferinz.
That Pneumatophilus Odhn., and Leptophallus Liihe, are to be
included along with these, as Odhner has indicated (2. p. 56),
appears to me somewhat doubtful.

The third form which I have to describe here is one of very
great interest. It was obtained from a Diamond Water-snake
(Tropidonotus rhombifer) from North America. The habitat,
unfortunately, is not recorded. It bears a certain resemblance to
the foregoing species, and belongs to the family Lepodermatide,
but it possesses an individuality sufficiently marked to constitute
a distinct generic type.

DasYMETRA CONFERTA, gen. et sp.n. (Plate XXVIII. figs. 8-10.)

The collection consisted of about a dozen specimens, all of which
were mature, and measured 35-46 mm. in length. The body is
elongated, slightly flattened and of fairly uniform breadth. The
greatest breadth occurs about the middle and is 1-1-4 mm. The
length is therefore about 34 times the breadth. The cuticle is
beset throughout its whole extent by long straight spines. It
appears to be somewhat deciduous, and in many specimens is
absent from a considerable part of the body, especially towards
the posterior end. Several specimens, however, retained the
cuticle and spines quite intact.

In a specimen of average length (4:2 mm.) the oral sucker
measures °56 mm. in diameter. It is globular, almost terminal
and not very muscular. The ventral sucker is practically of equal
size, if anything a trifle less. It is somewhat transversely oval,
the dimensions being 52 x-57 mm, It is only slightly prominent,

684 DR. WILLIAM NICOLL ON

not very muscular, and is situated 1-7 mm. from the anterior end.
The neck, therefore, comprises 2 of the body-length.

The alimentary canal is highly developed. It consists of a very
short prepharynx, with an enormous pharynx measuring *28 mm.
in diameter. The esophagus is shorter than the pharynx, being
only about -2 mm. long. It is fairly wide, with well-developed
musculature and numerous peri-cesophageal cells. It divides into
two very wide diverticula, which extend along the sides of the
body to near the posterior end. From the latter they are
separated by a loop of the uterus. The ends are slightly inflated
and somewhat turned in.

The excretory system has the same general structure as in the
two previous species. The main stem of the excretory vesicle
divides close behind the sheli-gland into two limbs, which extend
a short distance in front of the ventral sucker. From the vesicle
numerous twigs are given off, which divide and subdivide in the
lateral fields to form an intricate anastomosis. ‘The most peculiar
feature of the excretory system, however, is the pigmented con-
dition of the excretory tubules, which renders them strikingly
conspicuous and marks out their course with great distinctness.
This feature renders thespecies uniqueamongst the Lepodermatide.
The pigmentation is due to the excretory granules, which are
almost black in colour, and which fill the tubules. Only a few of
these are to be met with in the vesicle. A main excretory tubule
runs along the greater part of the length on each side of the body,
ventral to the intestinal diverticula. In front of the ventral
sucker it divides into small branches, one of which runs in to join
the vesicle, and another runs forward to the oral sucker. At the
posterior end it also divides into several branches. It is impossible
to say whether this pigmented condition occurs in life or is
a post-mortem appearance, but it was certainly present in all the
specimens.

The genital aperture is median, just over the intestinal bi-
furcation (‘3 mm. in front of the ventral sucker). It shows
a tendency to be deflected very slightly to the left side. In every
specimen the long, thick cirrus was exserted. The cirrus-pouch
is short and stout, in some cases being almost globular. Its
posterior end lies dorsal to the middle of the ventral sucker, but
it may extend beyond this to almost the posterior border of the
sucker. The pouch has an external wall composed of very thick
longitudinal muscle-fibres, with an inner layer of much smaller
circular fibres. It contains a small, slightly-coiled vesicula semi-
nalis, a small bulbous pars prostatica, with numerous prostatic
cells, and a long ductus and unarmed cirrus. As already men-
tioned, the latter was exserted im every case, so that the
arrangement depicted in fig. 8 (Pl. XX VIII.) must be regarded as
hypothetical. The vesicula, prostate, and cirrus all have a very
well-marked layer of longitudinal muscle-fibres,

The testes are situated obliquely, the left being well in front of
the right, but not entirely so. The former les about °3 mm., and
the latter ‘8mm, behind the ventral sucker. They are large ovoid

NEW TREMATODES FROM REPTILES. 685

bodies, with entire margins, and their long axes lie nearly in the
longitudinal axis of the body. They measure *5—7 mm. in length
and ‘4—"5 mm. in breadth. They are most remarkable, however,
for their great thickness, which is equal to or greater than the
breadth. They thus occupy nearly the whole body-thickness, a
fact which prevents them being obscured by the uterus. ‘They
are separated from each other by the wide ascending limb of the
uterus, against which they press, and their outer margins are
closely apposed to the intestinal diverticula.

The ovary is situated over the right posterior quadrant of the
ventral sucker, a short distance behind the end of the cirrus-
pouch. It projects well beyond the sucker, and is transversely
oval, measuring °35 mm. xX ‘28 mm. It lies close up to the dorsal
surface of the body. Close to the inner side of the ovary and
a little behind it, les a large shell-gland, with a short ootype.
A receptaculum seminis is absent, but Laurer’s canal is present.
The yolk-glands are rather voluminous. ‘They are entirely
lateral and peripheral, and they extend from the level of the
genital aperture, or rather behind it, to near the posterior border of
the right testis. Here, again, the anterior limit is fairly constant,
but the posterior limit is somewhat variable, less so, however, than
in the two species already described. The follicles are large and
arranged in regular dendritic groups. All the follicles are
connected up by short ducts. They le close under the surface
of the body, extending widely under the dorsal surface, but being
much more restricted ventrally, where they do not overlap the
inner wall of the intestinal diverticula.

The uterus is large and voluminous, but isnot much convoluted.
It is intermediate in type between that of Renifer and Ochetosoma,
but resembling the former more than the latter. It consists of
a small descending limb, which has a dorsal position and contains
a considerable number of ova. This runs to the posterior end of
the body where it forms a small convoluted knot, a fairly sym-
metrical pair of loops, one on each side, being thrown up towards the
ends of the intestinal diverticula. From this knot emerges the
ascending limb, which rapidly widens but does not attain its
maximum width till it has passed in front of the testes. 1t is
never so enormously dilated as in Lechriorchis validus. Near the
middle of the ventral sucker it runs into a remarkably powerful
vagina, which is as long as or somewhat longer than the cirrus-
pouch. The vagina possesses unusually muscular walls, there
being a very thick layer of longitudinal fibres and an equally
thick layer of circular fibres. It is surrounded by a great mass of
peri-vaginal cells, and it is lined by a thick layer of cuticle from
which small regular cuticular processes extend into the lumen.
The ova are numerous, dark brown and oval, with large well-
marked operculum. ‘They measure ‘033—'037 mm. in length and
‘016-019 mm. in breadth, the usual size being -036 x ‘018 mm.

The genus Dasymetra may be defined as follows :—
Lepodermatide; with moderately flattened body, entirely covered
Proc, Zoou. Sov.—1911, No. XLVIII. 48

686 ON NEW TREMATODES FROM REPTILES.

with spines. Intestine with large pharynx and wide diverticula
which extend near but not quite to the posterior end. Ex-
eretory vesicle Y-shaped with numerous side twigs. Genital
aperture median, a short distance in front of the ventral sucker.
Cirrus-pouch short and plump ; vesicula and pars prostatica short ;
cirrus long. Receptaculum seminis absent ; Laurer’s canal
present. Yolk-glands extensive, dendritic, peripheral. Uterus
forming a small convoluted knot at the posterior end of the
body, with a wide unconvoluted ascending hmb. Vagina long
and very muscular. Ova about ‘035 mm. long.

Type, D. conferta, sp. n.

The type-specimens of these species are deposited in the
Museum of the Royal College of Surgeons, London. _Co-types at
the Zoological Society’s Gardens.

References.

(1) M. Brawn, 1902.—Fascioliden der Vogel. Zoolog. Jahrbiicher ;
Abt. f. Syst. xvi. pp. 64-67.

(2) T. OpHner, 1910.—Nordostafrikanische Trematoden, grossten-
teils vom Weissen Nil. i. Fascioliden. Results of the
Swedish Zoological Expedition to Egypt and the White
Nile, 1901. No. 23 A, pp. 22-76.

(3) J. Poirier, 1886.—Trematodes nouveaux ou peu connus.
Bull. Soe. Philomat. Paris, sér. 7, vol. x. pp. 24-6.

(4) H.S. Prarr, 1903.—Descriptions of Four Distomes. Mark
Anniversary Volume, pp. 23-38.

(5) J. Starrorp, 1905.—Trematodes from Canadian Vertebrates.
Zool. Anzeiger, xxviii. p. 691.

EXPLANATION OF THE PLATES.

The following letters apply to all the figures :-—

D.St. Yolk-glands. | R.S.Ut. Receptaculum seminis
Hx. Excretory vesicle. | uterinum.
Ex.T. Excretory tubules. | T.,T;, T2. Testes.
J. Intestinal diverticula. | Ut. Uterus.
K.St. Ovary. | Vg. Vagina.
P.Pr. Pars prostatica. | V.S. Vesicula seminalis.

Prate XXVII.
Lechriorchis validus.
Fig. 1. Young specimen. Ventral view. X 25.
. Adult specimen. Ventral view. X 20.
. Transverse section near ends of intestinal diverticula. X 53.
. Transverse section, immediately in front of ventral sucker. X 50.
. Ovum. X 500.

OP cw be

Prats XXVIII.

Ochetosoma formosum.
. Ventral view. X 30.
. Ovum. X 550.
Dasymetra conferta.
. Ventral view. x 30.
. Transverse section, a little in front of ventral sucker. X 60.
, Ovum. X 5600,

ics!

Hes

oy .
ee
SOM WOH

Vee See nine WEALD ODS

H.Goodehild del et hth. Huth imp.
THE CHINESE TAKIN,
IBVUNSONRUCYATS) IEE IDE ORY.

ON MAMMALS FROM CENTRAL CHINA, 687

32. The Duke of Bedford’s Zoological Exploration of astern
Asia.—XIV. On Mammals from Southern Shen-si,
Central China. By Otprimip THomas, F.R.S., F.Z.S.*

[Received and Read April 25, 1911.]
(Plate X XTX. f)

As already indicated in paper No. XIII. of the present series +,
Mr. Malcolm Anderson and his party, working on behalf of the
Duke of Bedford, obtained a considerable number of mammals in
Southern Shen-si before going on to Kan-su and Sze-chwan,
whence the series described in that paper was collected. Owing
to delay in transport, however, a large part of the Shen-si
collection has only recently arrived, and the diagnoses of J/yotis
myosotis ancilla, Microtus nux and M. johannes § are all that have
been published upon it.

The present paper gives a list of all the specimens obtained in
8. Shen-si by Mr. Anderson between his second landing in China
in the autumn of 1909 and his move on into Kansu in 1910.

The regions explored were, firstly, the district round Shang-
chou, 8.E. Shen-si (about 33° 40’ N., 110° 20' E.), and, secondly,
the important mountain Tai-pei-san (about 34° N., 107° 30’ E.),
one of the sacred Chinese mountains, another being Omi-san,
Sze-chwan, where at a later period Mr. Anderson obtained the
many new species described in my Sze-chwan paper.

Of the majority of the Shen-si specimens there is little new to
record, as Mr. Anderson had obtained the same species on
his previous visit to the more northern part of the province. But
in any case their interest is dwarfed by the discovery on Tai-pei-
san of a magnificent species of Takin, quite different from the
known W. Chinese species Ludorcas tibetanus, and both in interest
and beauty one of the most striking {mammals that it has ever
been my good fortune to describe.

Besides this fine animal, of which a coloured figure is given
(Pl. XXTX.), Mr. Anderson obtained a new Badger, a new Pika,

and a new Vole ||.

1. RHINOLOPHUS FERRUM-EQUINUM Schreb.
@. 2079, 2080, 2081. Shang-chou District, S.E. Shen-si.

* Published by permission of the Trustees of the British Museum.
+ For explanation of the Plate see p. 695.
P. Z. S. 1911, p. 158.

» P.Z.S. 1910, p. 635.
|| The complete account of these new forms appears here, but the names and

preliminary diagnoses of the species underlined were published in the ‘ Abstract,
No. 95, 1911.—Epitor,
48*

Crt

688 MR. OLDFIELD THOMAS ON

9. Myoris MYOSOTIS ANCILLA Thos.

3. 2082, 2083, 2084. 9. 2085. Shang-chou Dist., S.H.
Shen-si.

The typical series (cf. P.Z.S. 1910, p. 636), no. 2082 (B.M.
No. 10.5,.2.4.) the type.

3. CrocipuRA ATTENUATA M.-Edw.

g. 2009. King-tze-Kwan, 8.W. Honan. 850’.

4, CrocipurA coREx Thos.
3g. 2185. 30 miles S. of Feng-hsiang-fu, 8. Shen-si. 7000’.

5. FELIS FONTANIERI M.-Edw.

2. 2038. Shang-chou District, 8.E. Shen-si. 3000’.
This fine Leopard is a valuable accession to the Museum
collections.

6. Fexis microtis M.-Edw.
$. 2176. 30 miles 8. of Feng-hsiang-fu, 8. Shen-si. 3600’.
7. VIVERRA ZIBETHA, subsp. ?

9192. Native skin. 40 miles N. of Han-chung-fu, Shen-si.

This specimen does not agree with the description of the Civet
called V. filehneri by Matschie, but how far the colour-characters
used by him are likely to be diagnostic in so variable a group I
am not at present prepared to say.

8. PAGUMA LARVATA Gray.

6. 2053. Shang-chou Dist., S.E. Shen-si. 2300’.

9. VULPES sp.

3. 2178. 30 miles S. of Feng-hsiang-fu, 8. Shen-si. 3600.
10. Lurreoa srpirica Pall.

3s. 2070. Shang-chou Dist., 5.E. Shen-si.

@. 2118. Si-ngan-fu, Shen-si. 1200’.

11. ARCTONYX LEUCOLEMUS ORESTES.

Thos. Abstr. P. Z.8. 1911, p. 27 (May 2).

Q@. 2191 (young adult). Tsin-ling Mts., 34° N., 107° 45' E.,
S.W.Shen-si. Alt.12,000'. 25 January,1910. B.M. No.11.6.1.6.
Type.

Distinguished from the true lewcolemus of Peking by the
following characters :—

Dark mark enclosing eye not broadly projected forwards and
downwards to the upper lip, but practically confined to a strong
spectacle-mark, barely half an inch broad, surrounding the eye ;
a narrow and indistinct line only running forwards to the base
of the whiskers, the upper lips quite white. Dark patch behind
mouth much reduced, a trace only of it running forward to the

MAMMALS FROM CENTRAL CHINA. 689

angle of the mouth, separated from its fellow of the opposite side
by a white interramial space over an inch broad. Light throat-
patch not really white, but brownish white, figured in lewcolemus
as snowy white. Light patch in front of ear less prominent,
continuous with but darker than that under the eye. White
ear-rim much broader and more prominent. Back more broadly
washed with whitish than appears to be the case in lewcolemus,
the dorsal hairs white for their terminal 15-20 mm., while only
their points are said to be white in lewcolemus. Tail wholly
white.

Dimensions of the type, measured in the flesh :—

Head and body 570 mm.; tail 195; hind foot 94; ear 45.

Skull: condylo-basal length 132 mm. ; basal length 123; greatest
breadth 72; interorbital breadth 29; palatal length 86 ; greatest
diameter of m’* 16,

Hab. and type as above.

The British Museum had previously possessed no examples of
the N. Chinese Arcéonyx, so that this fine specimen is a valuable
accession. On account of the differences above detailed I cannot
refer it to the true lewcolemus, but think it represents a special
subspecies, as is the case with so many other Shen-si mammals.

12, PETAURISTA ALBORUFUS * M.-Edw.
2194. Native skin. Near Pao-Ning-Fu, N. Sze-chwan.

13. Trocoprerus xANTHIPES M.-Edw.
2055. Shang-chou Dist., S.E. Shen-si.

14. SciuROTAMIAS DAVIDIANUS M.-Edw.

3g. 2022, 2037, 2052, 2065, 2066, 2067, 2069, 2087, 2095,
2104. 9. 2054, 2059, 2096, 2105. Shang-chou Dist., 5.H.
Shen-si.

3. 2106, 2115. Ching-ling Mts., Lo-nan-hsien, S. Shen-si.

@. 2107. Ching-ling Mts., Lo-nan-hsien, 8. Shen-si.

15. EPIMys CONFUCIANUS, subsp.

3. 2034, 2063, 2064. 9. 2024, 2033, 2035, 2036, 2039.
Shang-chou Dist., S.E. Shen-si.

6. 2114. Ching-ling Mts., Lo-nan-hsien, 8. Shen-si.

@. 2010. King-tze-Kwan, 8.W. Honan.

S. 2012,2014. 2. 2015,2016. Shan-nan-hsien, 8.E. Shen-si.

So. 2122, 2123, 2137, 2140, 2143, 2144, 2152, 2161. 2. 2162.
30 miles 8. of Feng-hsiang-fu. 3600’.

In my previous paper the Kan-su specimens of this group were
assigned to #. confucianus luticolor, though tending to intergrade
with the typical H. confucianus of Sze-chwan. But now, on
laying out the whole of the N. China representatives of the group,

* Petaurista is masculine, this being one of the numerous names which by their
deceptive form render desirable the suggested convention that all generic names in
zoology should be treated as masculine. See Stebbing, ‘ Knowledge,’ xxxiii. p. 259,
1910.

690 MR. OLDFIELD THOMAS ON

I find that the true Z. ¢. luticolor is a pale desert form peculiar
to the region towards the Ordos desert, that the present 8.
Shen-si specimens are, as is geographically correct, intermediates
between the /. c. sacer of Shantung on the east and the Kan-su
form on the west, and that the latter is sufficiently differentiated to

have a special subspecific name of its own. It may be called —

HPIMYs CONFUCIANUS CANORUS, subsp. n.

General colour approximating to “ clay-colour,” slightly darker
and more tawny than in sacer, much darker than in Jluticolor,
lighter than in confuctanus. Median darker dorsal line averaging
more distinct than in confucianus, less than in sacer. White of
under surface more markedly tinged with buffy or cream-colour
than in the other subspecies (but there is a doubt as to how much
this fades after death). Metatarsals without, or with but slightly
marked, darker patches. Tail with the dark colour of its upper
surface passing nearly or quite continuously to the end; its tip
hardly so heavily tufted as in sacer.

Skull about as in confucianus and luticolor, smaller than in
sacer.

Dimensions of the type, measured in the flesh :—

Head and body 120 mm.; tail 180; hind foot 27; ear 21°5.

Skull: greatest length 34°5 mm.: condylo-incisive length 30:5 ;
upper molar series 6.

Hab. Southern Kan-su, grading eastwards into sacer and south-
wards into true confucianus. Type from Wen-hsien Country,
S. Kan-su.

Type. Adult female. B.M. No, 11.2.1.110. Original number
2282. Collected 6 May, 1910.

16. Mus waGnert Eversm.
3. 2094. Shang-chou Dist., S.E. Shen-si.

17. APODEMUS SPECIOSUS PENINSUL& Thos.

3. 2046, 2076, 2077. ©. 2045, 2047, 2092, 2093, 2103.
Shang-chou Dist., S.H. Shen-si.

3. 2113. Ching-ling Mts., Lo-nan-hsien, 8. Shen-si.

Q. 2126, 2132, 2136. 30 milesS. of Feng-hsiang-fu, 8. Shen-si.

3. 2167, 2169, 2172. 9. 2166, 2168, 2174. Tai-pei-san.
10,600’.

18. APODEMUS AGRARIUS PALLIDIOR Thos.

3. 2030. 2. 2031, 2040, 2057, 2058, 2061. Shang-chou
Dist., S.E. Shen-si.

3. 2019. 9. 2020. Shan-nan-hsien, S.E. Shen-si.

@. 2108, 2109, 2112. Ching-ling Mts., Lo-nan-hsien, 8.
Shen-si.

96,99. 30 miles S. of Feng-siang-fu, S. Shen-si. 3600’.

19. Micromys minutus M.-Edw.
3. 2056. Shang-chou Dist., S.E. Shen-si.

MAMMALS FROM CENTRAL CHINA. 691

20. CriceruLus Trrron Thos.

@. 2011. King-tze-Kwan, S.W. Honan.
3. 2013, 2018. ‘Shan-nan-hsien, 8.E. Shen-si.
2. 2028, 2029, 2062. Shang-chou Dist., S.E. Shen-si.

21. CRICETULUS ANDERSONI Thos.
3S. 2116, 2117. Si-ngan-fu, 8. Shen-si. 1200’.

22. MicroTus CALAMORUM SUPERUS.
Thos. Abstr. P. Z.S. 1911, p. 27 (May 2).

6. 2121, 2127, 2128, 2129, 2138, 2139, 2147, 2154, 2163,
2181, 2182, 2183. .

On 2119, 2120, 2130, 2146, 2155, 2164, 2179, 2180. 30 miles
8. of Feng-hsiang-fu, S.Shen-si. 3600’.

A larger longer-tailed race of the Nanking Vole.

Fur longer and finer than in true calamorwm, the hairs of the
back about 16 instead of 10 mm. in length. Colour quite as in
calamorum, except that all the specimens have a dull buffy suffusion
in the surface-colour of the abdomen, this being only the case in
certain of the younger examples of calamorum, the older ones
having the belly clear greyish white. Tail longer than in cala-
morum, the usual length in adult specimens 60-63 mm., as against
52-53 in that animal, its coloration more prominently bicolor.

Skull like that of calamorwm, but slightly longer than in speci-
mens of similar age. Nasals longer. Bullw generally larger.

Dimensions of the type, measured in the flesh :—

Head and body 130 mm.; tail 63; hind foot 24; ear 13.

Skull : condylo- -basal lensth 33 mm. ; condylo-incisive length
33°2; zygomatic breadth 17:8 ; nasals 8°5 x 3°6.

Type. Adult male. B.M. No. 11.6.1.45. Original number
2163. Collected 7 January, 1910.

This Shen-si representative of the Lower Yang-tze Reed-Vole
is distinguishable by its longer tail and longer fur, the latter in
obvious correlation with the greater altitude at which it is found.

23. MIcROTUS MANDARINUS M.-Edw.

2. 2032. Shang-chou Dist., 8.E. Shen-si.
Skull. S.E. Shen-si.

The specimens referred to in the description of J/icrotus
johannes (P. Z.8. 1910, p. 637).

24. Microtus (CARyomys) Nux Thos.

3. 2041, 2042, 2043, 2072, 2073, 2088, 2089, 2090, 2097,
2098. 2. 2044, 2050, 2091, 2100. Shang-chou Dist., S.E.
Shen-si.

3. 2110, 2111. Ching-ling Mts., Lo-nan-hsien, S. Shen-si.

Described from these specimens, P. Z.5. 1910, p. 636. No.
2089 (B.M. No. 10.5.2.79) the type.

692 MR. OLDFIELD THOMAS ON

25. Microrus (Caryomys) EvA Thos.

3S. 2173. Tai-pei-san, Tsin-ling Mts., S.W. Shen-si. 10,600’.

The typical series was obtained near Tati-chow, Kan-su, at a
similar elevation.

26. Lepus swinHoE! Thos.

3. 2145. 30 miles 8. of Feng-hsiang-fu, 8. Shen-si. 3600’.

Represents L. filchneri Matsch., and also the Tai-pei-san
subspecies “ suppressed ” by Dr. Allen*, to which he, nevertheless,
attaches the name L. swinhoer brevinasus.

Of all the many examples of ZL. swinhoei obtained by
Mr. Anderson, whether topotypes from Shantung or from other
localities, this specimen is one of those that agree most closely
with the type collected by Swinhoe, owing to its being a dark-
coloured individual, with its general buffy coloration more or less
suffused with pinkish.

27. OCHOTONA SYRINX.

Thos. Abstr. P.Z.S8. 1911, p. 27 (May 2).

g. 2170. @. 2171. Tai-pei-san. 10,600’.

(2) S. 2071 (young). Shang-chou Dist., 8.E. Shen-si. 3300’.

Related to O. cansa, but larger. Bulle smaller.

Size distinctly larger than in O. cansa and O. sorella. Fur (in
winter pelage) long and soft, not very thick ; hairs of back about
16-17 mm. in length. General colour more blue-grey throughout
than in cansa; head grey tinged with clay-colour ; nape ‘‘smoke-
grey”; dorsal area darker grey tinged with brown ; rump nearly
“mouse-erey.” Flanks near ‘“broccoli-brown.” Under surface
greyish, the bases of the hairs slaty, their tips greyish white.
Ears greyish brown, their proectote blackish, their rims white ;
the tuft of long hairs at their inner base nearly “ cinnamon.”
Hands and feet white above, the thickly haired palms and soles
‘“‘ smoke-grey.”

Skull of the same general type as in O. cansa, of similar flat-
tened form, the palatal foramina not subdivided. Size distinctly
greater throughout ; nasals of about the same shape, not narrowed
as in QO. sorella. Interorbital space and brain-case broader.
Bulle conspicuously smaller or at least lower, those of O. cansa
being very large for so small a species.

Dimensions of the type, measured in the flesh :—

Head and body 142 mm.; hind foot 28:5; ear 16.

Skull: greatest length 36°5 mm. ; condylo-incisive length 33°5 ;
zygomatic breadth 18°5; nasals 11°8x 5-4; interorbital breadth
4-3; breadth of brain-case 15:7; palatilar length 12; palatal
foramina 9°2 x 3:7 ; upper cheek-tooth series (alveoli) 7:1.

Hab, Tai-pei-san.

* Bull. Am. Mus. N. H. xxvi. p. 427, 1909.

MAMMALS FROM CENTRAL CHINA. 693

Type. Adult male. B.M. No. 11.6.1.59. Original number
2170. Collected 9 January, 1910.

This is probably the species recorded as O. cansa by Dr. Allen *,
who had, however, only “skins with fragmentary skulls” for
comparison with the single type of Dr. Lyon’s species. The
present perfect specimens, compared with the topotypical series

of O. cansa obtained in Kan-su by Mr. Anderson, show that the
' species is readily distinguishable, both by its greater general size
and its much less swollen bulle.

No. 2071 is too young for certain determination, but it is
interesting as still retaining in place the minute milk-predecessors
of the small posterior incisors. The milk-teeth are pressed close
against the front face of the permanent teeth, between these and
the large incisors in front of them.

28. Sus sp.
@. 2186. 30 miles S. of Feng-hsiang-fu, 8. Shen-si.

In the state of confusion to which the late Pere Heude
reduced the systematic arrangement of the Chinese members of
Sus, as with every other genus with which he dealt at any length,
it is impossible to give a satisfactory determination of this Wild
Boar at present. It probably represents the Sus oxyodontus of
Heude, from the Upper Han, but whether that is or is not a valid
species I am unable to express an opinion.

29. Buporcas BEDFoRDI. (Plate X XIX.)
Thos. Abstr. P. Z.8. 1911, p. 27 (May 2).
3. 2175. 9. 2189, 2190. Tai-pei-san. 10,000’.

A wholly pale buffy species, practically without darker
markings.

Fur longer and richer than in the available specimens of B. tebet-
anus, but the dates of these are not known, while the examples of
B. bedfordi were killed in mid-winter; hairs of sides of neck
attaining 7-8 inches, and those of back 3-4. General colour a
beautiful glossy golden-buffy, more cream-buff in the females, more
tending towards ochraceous in the male; at least at the ends of
the hairs, their bases being still creamy or whitish. Darker
markings on muzzle, ears, hinder back, and limbs, characteristic
of B. tibetanus, practically absent. No. 2190 (2) entirely without
darker hairs on the muzzle, 2175 (¢) with a few, and 2189 (9?)
with more, but the imeonspicuous darker patch so formed very
different from the large and prominent black mask characteristic of
B. tibetanus. Dorsal line not darkened at all, its hairs elongated
and more strongly buffy in the male, not differentiated at all in
the females. Limbs not obviously darkened terminally, though a
few isolated darker hairs are present on the carpus and tarsus.

‘ * Bull. Am. Mus. N. H. xxvi. p. 427, 1909.

694 MR. OLDFIELD THOMAS ON

Tail bushy, a few darker hairs on it in No. 2189, soiled ochraceous
buffy in the others.

Skull apparently rather smaller, and its nasal region less vaulted
than in either tawicolor or tibetanus, but, owing to differences in
the ages of the specimens available, a satisfactory comparison 1s
not at present possible.

Dimensions of the male, measured by Mr. Anderson in the
flesh :—

Head and body 1925 mm. ; tail 23; hind foot 340; ear 127.

Skulls :—

3 2
(young adult). (adult).

Condylo-basal length ....................0006 389mm. 381 mm.
Ly Soma tich Ocead ble sne eae. oseee tae eere 176 164
Height of nasal convexity above middle

OL OL NCTC reer a Nan, arn ennm AR Ion Ltos 118 113
Length of muzzle, to front of p*......... 113 115
Palatal Mlemothim steers eee cere eran cere ter 245 24)
Greatest spread of horns, on outer edge... 401 308

Type. Adult female. B.M. No. 11.6.1.64. Original number
2190. Killed 15 January, 1910.

The discovery of this splendid animal, whose golden-bufty colour
renders it by far the most beautiful of its genus, is of the highest
interest, and it is with great pleasure that I name the species in
honour of the Society’s President, during whose exploration of
Eastern Asia it has been obtained. Mr. Anderson himself seems
to have thought the occurrence of Takin on Tai-pei-san of special
interest, and believed that they would probably prove to be new.
He says: ‘‘ The herds on Tai-pei-san are isolated by some hundreds
of miles from the nearest others we could hear of, and as I could
not learn that any other foreigner has hunted them on Tai-pei, I
believe the chance for a new species is good.”

As a matter of fact, however, specimens had previously been
obtained and had passed into the possession of the American
Museum of Natural History at New York. But these were quite
young, and showed, as it was not unnatural that the young should
show, more or less of the normal coloration of the group, with
blackish muzzle and extremities, and therefore in recording them
Dr. Allen* saw no reason to suppose them different from
B. tibetanus. The practically unicolor condition of b. hedfordi
proves therefore to be a characteristic of the adult, a fact which,
in view of the peculiar specialization of such a colour, is not at
all surprising.

Even in B. tibetanus, as shown by Milne-Edwards’s figure ft, the
young is very materially darker than the adult.

That the Takin of Tai-pei-san and other parts of the Pe-ling
range should be different from that of the mountains of Sze-chwan

* Bull. Am. Mus. N. H. xxvi. p. 4256, 1909.
+ Rech. Mamm. Atl. pl. 74.

MAMMALS FROM CENTRAL CHINA. 695

is only in accordance with the indications furnished by the
remainder of Mr. Anderson’s collections, as brought out in the
paper No. XIII. of the present series *.

With regard to other names that have been given in the genus
Budorcas, Mr. Lydekkert has shown that both sinensis and mitchella
are synonymous with tbetanus, so that there appears to be no
existing name which comes into question in now describing this
beautiful species.

It may be noted that on each side of the withers of the two
females there isa large patch of grey hairs, these hairs being horny
whitish, curiously ringed or beaded with black. Similar hairs
have been found on a specimen of B. tibetanus, and so would
appear to be natural and not discoloured artificially as I at first
supposed. No such patch is present on the male.

Mr. Anderson has sent me the following further notes on this
animal :—

“'Takin.—Found in large herds on Tai-pei-san, where it lives
in precipitous places at from 9,000 to 11,000 ft. altitude. Its
food is bamboo grass; a small bamboo very common at these
altitudes. Although apparently clumsy, I found these animals
very alert, and capable of picking their way very deftly and
rapidly down steep mountain sides. A herd my companions
and I saw seemed to contain about 40 individuals ; Chinese
hunters reported another herd of 80. These herds seem quite
isolated on the peaks of Tai-pei-san. Travelling westward we
did not hear of any Takins again till we reached Pie-kou in
Southern Kansu, and I am in doubt whether this was the same
animal or not.

Chinese name :—Pan-yang; at Tai-pei. The species in Sze-
chwan is called Yei-nu=Wild Cow. The name Pan-yang is also
applied to a mountain sheep or goat.”

30. NEMORHADUS sp.
@. 2188. 30 miles 8. of Feng-hsiang-fu, S. Shen-si. 10,000’.
Near the Long-tailed Goral (1. caudatus M.-Kdw.).

31. CAPREOLUS BEDFORDI Thos.
3. 2153 (young). 2. 2177, 2187. 30 milesS. of Feng-hsiang-
ily BOOP.

EXPLANATION OF PLATE XXIX.
The Chinese Takin (Budorcas bedfordi). Male.

* P.Z.S. 1911, p. 158. + P. Z.S. 1908, p. 795.

696 LT.-COL. N. MANDERS ON THE

: EXHIBITIONS AND NOTICES.
May 9, 1911.

E. G. B. Meapse-Wa po, Esq., Vice-President,
in the Chair.

My. R. I. Pocock, F.R.S., F.L.8., Superintendent of the Gardens,
exhibited some of the hair of the “puppy coat” of a Grey Seal
(Halicherus grypus), which was caught at Barmouth, in Merioneth-
shire, at the end of April. When received at the Gardens at
the beginning of May this Seal was covered, with exception of
the head and flippers, with longish woolly white hair, the last
of which was moulted on May 7th. Most authorities state that
Grey Seals are born in the autumn, not later than about the
middle of October, and that the puppy coat is shed from a month
to six weeks later. Allowing six weeks for the retention of its
puppy coat, this Welsh Grey Seal must have been born near the
middle of March, a date in tolerably close agreement with the
date, namely the end of February, given by Cneiff for the birth
of these Seals in the Gulf of Bothnia. It is, therefore, quite clear
that these Seals breed both in the early spring and the autumn.

My. E. G. BouLencer exhibited some living male specimens of
the Midwife Toad (Alytes obstetricans) carrying the eggs. He
also exhibited a number of the detached eggs to show the manner
in which they were strung together.

Mr. A. E. Anprrson exhibited a large number of photographs
of the more important fossil mammals in the Department of
Vertebrate Paleontology of the American Museum of Natural
History, New York, showing the methods of mounting fossil
skeletons. For comparison, a set of photographs was exhibited
with the skeleton supports eliminated from view, thus adding to
the pictorial value of pose in the specimens.

PAPERS.

33. An Investigation into the Validity of Miillerian and
other forms of Mimicry, with special reference to the
Islands of Bourbon, Mauritius, and Ceylon. By NEVILLE
Manpers, Lieut.-Colonel, R.A.M.C., F.Z.S., F.E.S.

[ Received May 8, 1911: Read May 9, 1911.]

Naturalists generally and the majority of entomologists are
disposed to accept a broad view that mimicry is caused by

VALIDITY OF SOME FORMS OF MIMICRY. 697

natural selection, but beyond this some are not prepared to go.
Others believe more or less implicitly m cryptic mimicry, either
active or passive, and that peculiar form of it known as Batesian
mimicry, that is, the resemblance for protection of a palatable to
an unpalatable species. And there are others again who, believing
in these, consider that Miillerian mimicry, that is, the resemblance
of unpalatable species for mutual protection brought about by the
tasting experiments of young reptiles and birds, plays a very
large part, even an overwhelming one, in the production of
mimicry.

The theories of the two great naturalists Bates and Miller
have now been before us for a great number of years ; but both,
the latter more especially, base their claims to recognition on
indirect evidence and not on experimentsand investigation in the
field. The exponents of these two theories maintain that though
direct evidence is largely absent, yet on no other reasonable
hypothesis can these remarkable cases of mimicry be explained.
The opponents, on the other hand, hold the view that as direct
evidence is possible though admittedly difficult to obtain, it ought
to be produced before either theory can be admitted as proved,
and until it is forthcoming they remain either actively hostile or
passively sceptical. It was with the feeling that both the sup-
porters of these theories and those opposed to them were equally
desirous of reaching some finality in this vexed question, which has
now been before them in one form or other for half a century—a
question which, with its periodical exhibition of violent eruption
and deceptive quiescence much resembles an Iceland geyser—that
I have during the past five years devoted as much of my time as
was practicable to the study of insectivorous birds and reptiles ;
and I have taken as my areas of investigation the islands of
Bourbon, Mauritius, and Ceylon, because the question is less
complicated on an island of small or moderate dimensions than on
such an extensive area as Africa or South America.

Before I left England in 1908 I had the great advantage
of being taken over the National Collection of Butterflies by
Mr. Guy Marshall, who, with his unrivalled knowledge of the
subject and persuasive powers, almost then and there made me
throw in my lot with the supporters of the Miillerian theory, but
in the midst of his arguments recollections of scenes in tropical
jungles obtruded themselves, and I was left in an irritating
condition of mingled belief and incredulity. A fairly extensive
reading of the whole subject consequently did little to convince
me, and my personal bearing towards both Batesian and Miillerian
mimicry was that the verdict must be the unsatisfactory one of
‘“‘not proven.”

I propose taking each of the islands in turn, enumerating the
reptiles and birds with their habits and distribution, directing
attention to the more striking cases of mimicry, and endeavouring
to ascertain on such data how far these theories are negatived or
sustained.

698 LT.-COL. N. MANDERS ON THE

Bourson.

Bourbon is a small circular island less than forty miles in
diameter, lying some three hundred miles from the east coast of
Madagascar. It has only twenty-two species of butterflies and
certainly one case of mimicry, which is very striking and quite
peculiar. It is that of a female Papilio (phorbanta) resembling
a Huplea. The group to whieh this Papilio belongs is green in
both sexes, but in this insect the female is dark brown and
resembles more or less closely the brown Huplea (goudoti)
occurring in the island. ‘There is no occasion to go into details,
as I have already brought them to the notice of the Entomolo-
gical Society in its Proceedings and Transactions, 1908, and have
figured both butterflies ; but I may add that they are essentially
insects of the littoral, common on one small portion of the coast,
particularly in gardens on the outskirts of. St. Denis, but very
rarely found above 1,500 feet or 2,000 feet. They are generally
associated.

Bourbon has no lizards with the exception of one introduced
species which is very rare. I was fortunate enough to find a
specimen; and I should say, judging by somewhat similar Ceylon
lizards, that in all probability butterflies would form part of its
diet, but it is far too rare to have any marked effect on the
butterfly population.

The following is a list of the insectivorous birds given to me by
a resident naturalist, which, so far as my knowledge extends, is
complete :—

1. The Sparrow. Passer domesticus.

. The Mynah. Acridotheres tristis.

. Zosterops (Malacirops) borbonica.

. Zosterops hesitata.

. Bec-Bee. Pratincola (Motacilla) sybilla.

. Coq des bois. Trochocercus borbonicus.

The Wheat Swallow. Phedina borbonica.

. The Little Grey-rumped Swiftlet. Collocalia francica.
. Le Merle cuisinier. Lalage (Oxynotus) newton.

© CO NIG) CU co

With regard to these, two, the Sparrow and Mynah, have been
introduced; on the former I need make no remark, it has the
same habits as its English relations. Two efforts have been made
in the last hundred years to instal the Mynah, but without success,
and this because it is considered a desirable morsel by the natives
and is mercilessly trapped and consequently very scarce; I saw
only one pair during my stay in the country.

Both species of Zosterops are very small birds, no bigger than
the English wren, and are found either singly or in small
family parties of five or six, flitting and creeping about the shrubs
after the manner of our long-tailed tits; they feed on nectar and
small insects. The Motacilla or Chat is of the same size and
much the same colouring as the Whinchat, and quite possibly feeds

VALIDITY OF SOME FORMS OF MIMICRY. 699

on the smaller butterflies, but would scarcely tackle an insect
considerably larger than our Swallowtail, but of this I have no
evidence. I saw it frequently in the gorge leading up to Salazie
and on the hills round St. Denis, but I saw nothing of it in the
neighbourhood of the town.

The Flycatcher (Z'rochocercus borbonicus) is the same species as
oceurs in Mauritius; it is said, on the authority of M.de Charmoy,
to feed on diptera and by preference on mosquitoes. It is
quite a small bird.

The Wheat Swallow (Phedina borbonica) has precisely the same
habits as regards its food as the English species ; it. appeared to
me to be fairly plentiful. It also occurs in Mauritius.

The Little Grey-1rumped Swiftlet (Collocalia francica) is the well-
known species that forms a nest of inspissated saliva. It occurs
in Mauritius. It is quite a small bird, decidedly smaller than our
Sand-Martin, and may occasionally snap up a small Lycenid.

Le Merle cuisinier, or Tui-tuit —Lalage (Oxynotus) newton.
I am unable to say whether this bird is abundant or not,
or whether it destroys butterflies. I did not come across it
in Bourbon, so I am inclined to doubt its being particularly
common. It has the same habits as the Mauritius ZLalage
rutiventer.

In the absence of any living bird it occurred to me that
possibly the extinct Bourbon Starling might have been the prime
factor in producing this case of mimicry, and I therefore wrote
to my venerable friend Dr. Jacob de Cortimoy for information
concerning it. | He is now verging on his ninetieth year, and is
probably the only one now living who has seen this bird alive ;
his letter is so interesting that I need make no apology for
transcribing a portion of it.

“T have known the bird you ask me about since childhood,
namely the Mregilupus varius (old writers called it F. capen-
sis), which has in fact entirely disappeared ...... When I was a
boy this bird lived in the forests of the interior of the island and
never set foot nor wing in towns or inhabited places. It rermained
faithful to the forests where it was bred, which it enlivened
with its clear notes. I used to hunt it then at an age when one
is pitiless. I can seeit now, a little larger than the white black-
bird, with a white crest on the head in the case of the male,
the wings a blackish grey on the upper surface, the beak and
feet yellowish. By no means shy, it was not frightened even by
the sound of firearms, and after a regular slaughter one went
off with dozens of these poor victims in one’s game-bag.

“After ten years spent in Paris I did not find a single one in
the forests where formerly they flew about in flocks. All ruthlessly
destroyed. I shall never forgive myself for the part, slight
though it was, which I took in the matter. I lost my taste for
sport and the best bag would not tempt me....... We will
now consider the feeding habits of this bird. Having raised
several in the aviary, I can risk talking about it though I never saw

700 LT.-COL. N. MANDERS ON THE

one feeding in the wild state. In my aviary its food consisted of
bananas, potatoes, and choux-choux, Sechiwm edule (boiled). But
when left to its own instincts, it must, like the other winged
denizens of the forest, have eaten insects as is done by its
companion in the forests, the Bourbon Blackbird (Hypszpetes
olivaceus) *, and as is the habit of most fruit-eating birds.”

This is a sad commentary on our boasted civilization, and I have
only to add that not half a dozen skins are now in existence.

This bird cannot have been in any way the cause of this
mimicry, as it inhabited the forest-covered hills in the interior of
the island, where these butterflies do not occur.

Papilio phorbanta female was figured by Boisduval in 1833 and
differs in no way from recent specimens. We may therefore infer
that the factor or factors which primarily induced this change of
colouring are still active ; but as there is no reptile or any bird
now living which attacks these butterflies as adults, it is difficult
to accept this as an effect produced by them. And it would
seem that the young of existing birds, with possibly one or two
exceptions, would be too small and feeble to attack these large
butterflies during their tasting experiments.

T now turn to the island of Mauritius, which lies some eighty
miles to the north of Bourbon and which is visible from there on
a clear day at certain seasons of the year.

MAURITIUS.

Mauritius has no arboreal lizards, and but one species of ground-
lizard, in appearance very like the English Sand-Lizard. It is
confined to the coast, and is I believe somewhat uncommon. I
found it in some numbers on the uninhabited islet of the [le de
la Passe at the entrance of Mahébourg harbour. It was quite
tame, even confidential, and made no display of timidity in taking
and eating a small piece of boiled potato presented to it on the
end of a fork. We may, I presume, regard this lizard as an
indiscriminate feeder !

My friend M. d’Emmerez de Charmoy, Director of the Port
Louis Museum, a Mauritius gentleman who has an unrivalled
knowledge of the fauna, and who has tracked, shot, skinned,
dissected and mounted the whole of the splendid collection of
Mauritius birds in the Port Louis Museum, has very kindly
favoured me with the following list and notes on the insectivorous
birds; it can be taken as complete, and I doubt whether any
local fauna of a tropical island is so completely known as is this
to M. de Charmoy.

T have added a few notes of my own in square brackets.

1. Le Mangeur de Poule (Tinnunculus punctatus) [| Cerchneis

punctata|.
Feeds preferably on insects rather than on small birds.

* This bird occurs in Mauritius, but was not given to me by the Curator of the
Museum as an insectivorous bird,

VALIDITY OF SOME FORMS OF MIMICRY. 701

I have many times discovered in their stomachs the remains
of locusts, field-crickets, and also stick insects.

[This Kestrel is rather smaller than the English bird and, like
it, is persecuted persistently. It may also feed on butterflies, but
it is so rare that its influence can be little felt; it is entirely
confined to the small portion of indigenous forest now remaining. |

2. Le Merle cuisinier (Oxynotus ferrugineus) |Lalage rufi-
venter. |

Ts essentially insectivorous; I have seen these birds catching
Mantis religiosa and I have found in their stomachs Scarabzei
(Cratopus) and fragments of moths’ wings.

[This bird of late years has become exceedingly rare and is
verging on extinction. On my telling M. de Charmoy that I
had seen a pair in the forest, he congratulated me with as much
fervour as if I had seen a Dodo!|

3. L’Oiseau Banane (Youdia erythrocephala).

Frequents very persistently bananas when in flower, and
captures the minute insects which are attracted by the honey of
these flowers; lives also on the petals of flowers and on small
lepidopterous larvee. |M. de Charmoy considers from his dis-
sections that this bird is incorrectly placed in this genus, which is
essentially a grain-feeding one. |

4. All these species are indigenous and so also are the two
species of Zosterops, Z. mauritiana and Z, chloronota, which live
almost entirely on the larve of lepidoptera. I am unable to give
the names of the kinds they capture, but no doubt they take any
kind of caterpillar.

5. Le Coq des bois (Muscipeta borbonica) | Trochocercus borboni-
cus|. Also indigenous; is an inhabitant of the forests and is
found also along river-courses; it chases diptera by preference
and particularly mosquitoes.

6. Le Boulbul (Pycnonotus jocosus) was introduced in 1892 by
M. Gabriel Reynard and is now to be found everywhere. It is
certainly to be found in great numbers, being often a plague. It
consumes the best fruits and vegetables and the blossoms of
fruit trees. I have often seen it hunting for moths, especially
for Ophiuside, and in the fields of wild indigo it captures Lyceenide.
[The most common butterfly in these fields is Lampides betica. |

7. Le Martin (Acridotheres tristis) was introduced from the
Coromandel coast by M. Boueher des Friyes, and by Pierre a
Mainard (2) into Réunion, to destroy the crickets which ravaged
the islands in 1759. It is found in great numbers in newly
tilled fields hunting after all sorts of insects, and especially after
the eggs of crickets.

The Fringillide cannot be considered insect hunters though
they catch one on the wing when they come across it. They
Proc, Zoon, Soc.—1911, No, XLIX, 49

702 LT.-COL. N. MANDERS ON THE

have not the slightest share in the reduction of local species (of
insects).

There are thirty species of butterflies, but with the exception
of Hypolimnas misippus and Danais chrysippus there is no such
well marked case of mimicry as that of the Euploea and Papilio
in Bourbon.

Professor Poulton has however thrown out the suggestion
that the female Papilio manlius, though green, is approaching
Euplea euphon, as it is of a distinctly brownish green tint
compared for instance with the Madagascar P. epiphorbas. _ Mr.
Trimen also considers that Danais (Amauris) phaedon and the
Euploea mimic each other to a certain extent. Instances of
seasonal dimorphism (cryptic defence) are however numerous.

The evidence I have collected does not seem to justify the
contention that the above instances of mimicry are due to the
depredations of old birds or to the experimental tasting of young
ones. Though no doubt experimental tasting of insects generally
takes place, it would appear that the birds are of such a character
that butterflies would not be attacked except to the smallest
extent.

CEYLON.

T now turn to the island of Ceylon. If the problem we are
considering has so far been simple owing to the small size of the
islands dealt with and their very limited fauna, it is by no means
so in Ceylon, which is far larger, being about two-thirds the size
of Ireland, with a wonderful diversity of hill and plain and equal
diversity of climate. Its general characteristics are too well
known to require repetition, and there are two hundred and sixty
species of butterflies.

>

Mimicry among Ceylon Butterflies.

Mimicry is by no means uncommon among Ceylon butterflies
and the following will serve as examples of it.

Mimic. MODEL.
Hypolimnas bolina 2. Huplea (several species).
rs misippus 2. Danais chrysippus.

Elymnias fraterna 9. » plexippus.
Argynnis hyperbius 2. © 55
Pareronia ceylonica @. » aglea.
Prioneris sita. Delias eucharis,
Papilio clytia race

lankeswara 3 Q. Huplea (several species).
Papilio clytia form dissimilis. Danais aglea (and allies).
Papilio polytes Q. Papilio aristolochie.

5s ., form romulus Q. Papilio hector.

And the Miillerian combination of the three Eupleeas, core, coreta,
and klugit.
Euplea coreta and £, core, asa reference to Mr. Moulton’s plate

VALIDITY OF SOME FORMS OF MIMICRY. 703

in Trans. Ent. Soc. Lond. for 1908 will show, are extremely alike,
yet under certain circumstances I can recognize them when on
the wing without great difficulty. When herded together in
shady jungle, as is their frequent custom, it is impossible to differ-
entiate them; but when flying singly over an open space, the
former appears a blacker and broader insect with a rather more
flapping flight, by which I can usually distinguish it from £. core.

The resemblance between Danais chrysippus and Hypolimnas
misippus 9 is well known, and I have often found them mixed
together in local collections, but on the wing under ordinary
circumstances differential diagnosis is by no means impracticable :
the female of the latter is almost invariably seen flying close to
the ground selecting favourable situations for oviposition, and
her method of flight, difficult to describe, differs from that of
D. chrysippus, which oviposits on a shrub some four or five feet
in height; when not thus engaged the two are undoubtedly
difficult to distinguish at about ten yards distance, and it is
curious how often the male of H. misippus mistakes D. chrysippus
for a female of its own species. Prioneris sita can at once be
distinguished from Delias eucharis by its rapid darting flight.

A ease of resemblance, though not always stated to be one of
mimicry, is that-of Argynnis hyperbius female and Danais plex-
ippus or D. chrysippus. This was remarked on by Butler so long
ago as 1884 and has been repeatedly noticed since, particularly
by Longstaff and Bainbrigge Fletcher. The resemblance is,
however, entirely accidental, as their habitat in 8S. India and
Ceylon clearly shows. Broadly speaking, if observed above 4000
feet, it will assuredly be 4. hyperbius, if on the littoral and up to
about 4000 feet, almost certainly D. plexippus; it is only on the
confines of each other’s territory that they in any way come in
contact and where an error can be made. Cethosia mnietneri,
Danais ceylonica, and Papilio clytia (dissimilis) fly in the same
localities, that is, in jungle where the light is very flickering. It
is not only very difficult to distinguish ‘them apart, but they are
quite difficult to see, as their black and white marking tends to
make them invisible. They also frequent the outskirts of jungle
and even more open country, and here they are quite easy to
distinguish, particularly P. clytia (dissimilis), which is a larger
and much yellower butterfly.

Mr. T. Bell considers Cethosia to be an unpalatable genus owing
to the nauseous juices and leathery bodies of the species. Huripus
consimilis very closely resembles a Danaid in both sexes, and
would be considered an undoubted case of either Batesian or
Miillerian mimicry: it is not a Ceylon butterfly, and I first
made its acquaintance in the Nilgiris; by its lofty sailing flight,
particularly in the female, I recognized it at a glance from a
Danaid, which rarely or never ascends more than about ten feet
from the ground.

It is more in their peculiar manner of flight rather than in
any difference of colouring that model and mimic can as a rule

4.9*

704 L.-COL. N. MANDERS ON THE

be distinguished; when at rest the difficulty is considerably
greater, and I would draw particular attention how not infre-
quently butterflies of a similar pattern on the under surface of
their wings chose the same resting places, and often the same
bush or branch on which to settle for the night. The following
two notes indicate this.

“ Colombo, July 1909. TI observed at sunset sixteen Zelchinia
viole and some Danais chrysippus with their wings closed, at rest
on a leafless bush; they exactly resembled withered leaves, and it
was exceeding difficult at ten measured paces to distinguish the
species, at about twenty paces it was very difficult to recognize
them as butterflies at all, and at thirty paces they were practically
invisible.”

“Kullar, Nilgiris, 12.4.1910. In a grove of Areca palms
Euplea coreta, £. core, Danais limniace and D, septentrionis simply
swarmed, they were in hundreds and hung in festoons from the
palms. Though many of each kind were on each leaf, they usually
kept together. It was deep shade, and the Euploas seemed to
match the decayed vegetation and the Danaids the green leaves.
There were also many Danais plewippus but no other butterflies.”

With regard to the above quoted instances, Mr. R. C. Punnett,
after a study of them for about six weeks, writes: ‘‘ With the
exception of Argynnis hyperbius and Prioneris sita, 1 have had
frequent opportunities of observing all these cases, and in every
one it has appeared to me that the resemblance is far less striking
when the insects are seen alive than when they are exhibited
pinned out in the orthodox way on cork. I have found that with
very little experience the eye comes to distinguish the mimic from
the model without hesitation. As a rule it is in the mode of flight
that they differ from one another...... My impressions of all
these so-called cases of mimicry which I have been able to see, is
that the resemblances are certainly not sufficiently close to
deceive the eye of a civilized man with a little experience of them.
For that reason I am inclined to doubt whether they would
systematically deceive an enemy brought up among them, whose
means of earning a livelihood depended largely upon the readiness
with which he could distinguish between mimic and model. I
do not wish to deny that in some cases, and upon occasion, the
resemblance may be of service.”

All local entomologists would endorse the above remarks, but
there is this to be said, that Mr. Punnett was aware of these cases
of mimicry before he arrived in Ceylon, and was on the look out
for them; if he had been totally ignorant of them, it would
have taken him some time before he recognized the phenomenon,
but having once done so, he would no doubt have had no further
difficulty.

The detection of certain cases of mimicry would appear to be
greater in some countries than in Ceylon. Colonel Bingham,
writing of the Papilio clytia group, says: “They have nearly all
4 wonderful resemblance to forms of Huplea and Danais, and it

VALIDITY OF SOME FORMS OF MIMICRY. 705

requires a quick eye and some experience to discriminate between
Euplea and Papilio clytia vace panope and between Danas
limniace and the dissimilis form of P. elytia, especially when on
the wing.” In reply to a query of mine on this point Mr, W. F. H.
Rosenberg writes :—‘ London, August 29th, 1910. During my
travels in Colombia and Ecuador I found that mimicking species
did frequent the same places as their models. For instance, the
Dismorphias (Pierines) which mimic certain groups of Ithomiine
butterflies, such as /thomia zelica, were seen flying in clearings
in woods ete., in company with the Ithomias. It is curious to
note, however, that there is a slight difference, difficult to
describe, in the mode of flight of the two groups, so much so
that a trained collector would rarely mistake one for the other.
Again, the S. American Acrvines of the genus dActinote, which
settle in groups on damp patches of sand, have their mimics in
the shape of Nymphalines of the genus Hresia. But while the
Actinotes will allow themselves to be picked up with the fingers,
the Eresias always fly up on the approach of danger.”

The Rev. St. Aubyn Rogers also, in his well known paper on
Kast African butterflies, mentions how often at first he was
deceived by model and mimie, but how after a short acquaintance
he readily recognized them.

But confining myself to Ceylon, the ease with which every case
of mimicry occurring there can usually be detected, causes me to
be in sympathy with those who consider that this constitutes a
great difficulty in the acceptance both of Batesian and Miillerian
mimicry. Both reptiles and birds are well represented, and in
order to afford a complete study of the fauna I propose taking
them in order, first dealing with the reptiles. I can scarcely
hope not to have overlooked some species, but I trust there is no
serious error.

The Lizards of Ceylon.

Five families ave represented; namely Geckonidee, Varanide,
Scincide, Lacertidee, and Agamide. The first three of these are
almost unquestionably of no account in our enquiry. The
Geckonide are mostly nocturnal with but one diurnal species,
Gonatodes kandianus, common in houses at Kandy and confined
to that part of the island. The Geckos I have observed feeding
at night were quite indiscriminate in their captures. Of the
Varanide, two species are large carnivorous lizards, commonly
called iguanas, frequently attaining a length of three or even four
feet. One species when young ascends trees, and Mr. Rosenberg
has seen a Mexican species eating butterflies.

The Scincide has four genera, Acontias, Chalcidoseps,
Lygosoma, and Mabuia. The first has four species with limbs
rudimentary or absent ; in appearance they are very like our slow-
worms but smaller. One species, 4. burtoni, is usually found
under stones, and the others, so far as I have been able to
ascertain, inhabit similar situations. They appear to feed on

g

“06. OP LT.-COL. N. MANDERS ON THE

small worms. Chalcidoseps has but one species, C. thwartesti, a
small creature about two inches long with very short limbs. It
is not represented in the Colombo Museum collection, and I am
unacquainted with it. Zygosoma has three species, one found in
the hills, the other two common in the low country. Their
forelegs are very feebly developed, in fact almost rudimentary,
and judging by their general appearance they probably have the
same habits as the next genus, Mabwia, which contains two
species, one of which is rather rare; the other, J/. carinaia, is the
well known Brahminy Scink, which so far as my experience goes
feeds almost entirely on ants, I have invariably failed to get it
to feed on butterflies.

The family Lacertidee has but one genus, Cabreta, containing a
single species leschenwultii, very small and rave, found only at
Mullative in the arid north-western district.

The Agamide has five genera, but three are represented by a
single species in each. Otocryptis hivittata, a very small creature,
confined apparently to the wet districts up to 2,000 feet: I have
no personal acquaintance with it. Cophotis zeylanica, also small
and found only in the hills: in captivity it feeds readily on flies,
which it captures after the well known manner of the chameleon.
Lyriocephalus scutatus, a magnificent creature found only in the
outer hill-ranges, where the climate is hot and moist. My friend
Mr. Alervs Hankey, who has kept these species both in captivity
and at large in his garden, informs me that they feed on “ almost
anything— moths, flies, beetles, grasshoppers, worms, and even
boiled vice.” We may conclude that their taste in butterflies,
which they in all probability eat somewhat extensively, is
impartial.

The genus Ceratophora has three species only, one of which I
know, C. stoddartii, found only in the bigher ranges. It feeds
on worms and positively declined butterflies when in captivity.

The genus Calotes has seven species, three of which are some-
what rare; these I do not know nor one other, C. mystaceus. Two,
C. ophiomachus and C. versicolor, are extremely abundant from the
coast to about 3,000 feet, thence upwards nigrilabris takes their
place and is likewise very common. All members of the genus
have probably the same habits and are undoubtedly highly
destructive to insect life. In fact I have little hesitation in
saying that they are the greatest enemies that butterflies have
to contend against, and when in Nuwara Eliya in 1909 I should
have had no great difficulty in making a fair collection of butter-
flies mutilated by them. The injuries were of all kinds, but
mostly a semi-circular piece, the size and shape of the lizard’s jaw,
had been taken out of the hind wing. These mutilated butterflies
indicated no discrimination on the part of their enemy ; perhaps
Argynnis hyperbius was the most frequent victim. My experi-
ments, though not so complete as might be wished, show that
these lizards exercise no partiality; but the butterflies experi-
mented with are those commonly found in the gardens at Colombo,

VALIDITY OF SOME FORMS OF MIMICRY. 707
where Calotes is abundant, and almost all those which occur at
Nuwara Eliya, the fauna of which is very poor.

Experiments with Lizards im Colombo.

The two species of reptiles experimented on belonged to the
genus Calotes (C. ophiomachus and C. versicolor), or blood-sucking
lizards as they are commonly called. In appearance and habits
they are far more like Chameleons than ordinary Sand-Lizards,
and like them have the faculty of changing colour and assuming
on occasions brilliant scarlet, yellow and green, particularly about
the head; but on the other hand they are very quick in their
movements and can run with considerable speed. They are
usually to be found sitting on walls and palings, clinging to the
stems of the longer grasses, and frequently lying in wait for their
prey behind the umbels of flowering shrubs, where their gaudy
colours help to deceive visiting insects. In such positions they
will wait motionless for hours on the chance of prey coming
within reach. But as I was unable to devote a whole day for
such prolonged investigations, I adopted the following method as
being the nearest approach to natural circumstances. I attached
a defunct or moribund butterfly by a long line of fine silk toa
fishing-rod, and thus succeeded with the help of the wind in
bringing the fly within reach of the reptile. The following were
the results from the end of November to the end of December
1908, with the notes I made at the time.

November 24. A male Papilio polytes was waved over a green
Calotes ophiomachus which was clinging to some grass. The lizard
became slightly excited and made a grab at the insect, biting out
a piece of the fore wing and immediately eating it; a second time
it bit a piece out of the hind wing including the tail; the insect
if alive would certainly have escaped on both occasions. Finally
it seized it by the head and ate the remainder.

November 30. A female Papilio polytes of the black and white
variety offered in the same way to another Calotes. This lizard,
which was evidently hungry, became excited when he saw the
butterfly, and made a grab at it and bit out a piece of the fore wing
which it promptly began to eat; the butterfly would have
escaped. While this “fishing” was going on, a male butterfly
attracted by the female flew quite close to the lizard, which made
a dart at it and tore away a piece of its wing, and the butterfly
flew off. A lizard grabs at any part of the butterfly that comes
within reach, and as the grass sways when it moves it is quite
a chance what part is seized; the habits of Calotes therefore
negative any theory of so-called directing marks so far as they are
concerned.

November 25. The wings of the black and white variety of
P. polytes ave quite common in the garden, and I am quite sure
the Calotes devour great numbers of them. I saw C. versicolor
with a live one in its mouth ; on this occasion it was holding the

708 Lt.-CoL. N. MANDERS ON THE

insect by the fore wing—when it saw me it ran to a tree, and [
fancy dropped the insect which, as I could not find it, probably
flew away.

December 4. Saw C. versicolor seize Delias eucharis on the
wing. On this occasion the lizard was in the foliage at the top
of a bush, and sprang out and captured the butterfly as it flew
past.

December 6. Huplaa core was eagerly seized, held in the
mouth for a long time and then eaten.

December 7. Telchinia viole was eagerly seized by the body and
devoured.

December 9. Two Danais chrysippus taken one after the other
by a green Calotes ophiomachus. It watched the butterfly, seized it
by the body, and ate the whole of it after holding it in its mouth
for some minutes.

December 18. Papilio (Menelaides) hector seized as soon as seen,
held in the mouth for some time and eaten slowly. The lizard
seemed very puzzled at the dryness of its meal as the butterfly
had been dead five days, but finally ate it all.

December 20. C. ophiomachus ate a male Papilio polytes.

December 30. Saw Terias hecabe captured by Calotes versicolor.

During these two months butterflies and other insects were very
numerous owing to the rain at the break of the N.E. monsoon.
The tall Mauritius grass in the garden was a place. of refuge for
the butterflies during the heavy rain, and numbers of them could
be seen any morning sunning themselves and sitting with
expanded wings halfway up the grass stems. /P. polytes and
P. demoleus were particularly numerous and several had pieces
taken out of then wings, no doubt by lizards. It was an
interesting fact that so long as the butterflies remained perfectly
still, they were entirely unnoticed by the lizards, though they
might be in close proximity to them. Both these butterflies
rest throughout the night with wings widely expanded.

Experiments with Lizards in Nuwara Eliya, 6,200 ft.

Three species were experimented on, Calotes nigrilabris, Cophotis
zeylanica, and Ceratophora stoddartii. All three are peculiar to
Ceylon and confined to the hill-districts. C. n2grilabris is about
a foot in length including the tail, and is brilliant emerald green
with a black bar across the lips. It has the same habits as
C. versicolor and C’. ophiomachus but is much tamer ; in fact it is
not at all difficult to capture with the hand as it rests on a bush.

Calotes negrilabris.

3.3.09. Ate a ZVerias hecabe greedily, and another shortly after.
3.3.09. The same lizard ate another 7’. hecabe, and another
made frantic grabs at H. core, dangled as usual at the end of a

VALIDITY OF SOMB FORMS OF MIMIORY. 709

string ; when eventually I allowed it to take it, it ate it readily.
The same lizard ate 7’. hecabe.

7.3.09. Another lizard ran out from its bush and caught
Terias libythea. A Papilio aristolochic, a very scarce species in
N’Eliya, caused great excitement in a male and female on the
same bush; they rushed out to capture it, then drew back
apparently frightened at its size and at length would have nothing
to do with it. Offered to another it was seized by the fore wing
and a part only eaten, the reason being that it was frightened of
me. Offered again to the first pair they took no notice of it, but
one made a jump of quite six inches and snapped up a fly which
had settled on a leaf. This shows that they were hungry at the
time.

15.3.09. A fresh Appias galene 9 readily taken, but not so
eagerly as by another which had recently changed its skin and to
which some of the slough was clinging. This was very hungry
and made quite a respectable jump at A. galene ¢ , which it caught
round the body. Immediately afterwards it devoured Catopsilia
pomona.

15.3.09. Offered Papilio (Menelaides) hector to a remarkably
fine lizard, which caught it by the base of the hind wings ; these it
ate very slowly and dropped the rest, no doubt because it was too
dry. The same thing happened with Velchinia viole and another
lizard. These two butterflies do not occur in N’Eliya.

16.3.09. A female moth, Spilosoma melanopsis, Fam. Arctiide,
with remarkably large brilliant crimson body and pink hind
wings, the dull fore wings being removed, was at once taken by
a lizard, which ran some distance after it. It proved a very
considerable mouthful which took quite half an hour to get rid of.

During March, April, and May I noticed a very considerable
number of butterflies, more particularly Argynnis hyperbius, with
pieces taken out of their wings, usually the posterior portion of
the secondaries. I have no doubt that almost all these injuries
were caused by this species of Calotes which is very numerous at
N’Eliya. I might almost say there is a specimen on every bush.

19.3.09. A Huplea core 3 proved very attractive; a half-
grown lizard ran more than a yard out of the hedge and seized it
by the head as it lay on the ground. With the same species a
large male made a dart at Danais fumata attached as usual to a
line of silk, caught it by the hind wing and devoured the whole of
it. A smaller individual seated on a bush of salvia became highly
excited by Polyommatus betica with its wings closed, and ran all
over the plant after 1t, eventually catching it by the body; imme-
diately afterwards it ate three Zerias hecabe, one after the other.

24.4.09. A female lizard,and one I have frequently experimented
on, ran from its hiding place and caught Pyrameis cardui by
the head after considering for a moment its cryptic underside.
Another scrambled over its bush in the usual way and seized
Lethe daretis 2 by the hind wing.

These lizards were very tenacious in their grasp: asa rule, after

710 LT.-COL. N. MANDERS ON THE

capturing their prey, they hold it quietly in the mouth for some
time, but if it struggles they immediately begin eating it. A
renewal of a struggle leads to the same thing.

Cophotis zeylanica.

This lizard is found only in the hill tracts of Ceylon, and is
most frequently found resting on tree-trunks in shady places.
It has considerable power of adapting its colouring to its
surroundings, but not to the same extent as the Chameleon. It is
almost five inches long including the tail, which is usually two or
three inches. It is extremely sluggish in its movements, depend-
ing entirely on its cryptic colouring to escape observation. It is
quite harmless and can easily be taken in the fingers. It eats flies
readily in captivity, but all my efforts, both with specimens at
large and in captivity, to induce them to eat butterflies resulted in
failure, even those with their wings entirely removed seemed only
to frighten them.

The Horned Lizard. Ceratophora stoddarti.

This interesting reptile is also peculiar to the island, entirely
confined to the hills, and I do not think it occurs below 5000 feet.
It varies in colour from a vivid green with black bands (three
only seen) to a uniform brown. It has.the faculty of changing
its colour according to the nature of its environment. Only
once have I found it otherwise than on a tree-trunk in deep shade,
where butterflies very rarely penetrate. All attempts to make it
eat butterflies in the wild state failed, and the following is a note
on a captured specimen :—‘“‘13.4.09. Put numerous flies in its
cage, there appeared. to be a diminution the following morning.
Put Verias hecabe and Pyrameis cardui alive into its cage. It did
not notice the latter as it rested with closed wings on the gravel,
- though within three inches of it; but when it moved its fore
wings up and down—not in and out—it was on the alert and
crept up to it, but before it had made up its cautious mind to
attack, the butterfly flew off, hitting it on the snout in so doing,
which utterly disconcerted it.” I may say that about the only
butterfly in the upper hill district which frequents shady places
is Lethe daretis. Mycalesis and Yphthima do not extend so high.
The chance of a meal off a butterfly is therefore remote.

Dr. Willey informs me that its natural food is small worms.

It would seem then that those who assume that reptiles take no
part in the production of Batesian or Miillerian mimicry are
correct, though further experiments are required.

The Insectivorous Birds of Ceylon.

A few introductory remarks are needed before dealing with the
insectivorous birds. Speaking generally the Mimicrists, if I may
use the term, maintain that birds do eat butterflies largely; the
Anti-Mimicrists that they do not. The evidence of the latter is

VALIDITY OF SOME FORMS OF MIMICRY. (Mal

necessarily negative, and we must therefore try to find some
reasonable standard by which we can judge whether a particular
bird is a butterfly eater or not. We shall not be far wrong if we
employ the criteria accepted by Mr. Guy Marshall in his paper
‘“‘ Birds as a factor in the production of Mimetic Resemblances
among Butterfiies” (Trans. Ent. Soc. Lond. 1909), only in this
case in a contrary sense. Speaking of the want of real evidence
on the part of the Anti-Mimicrists, he says:——“* When a naturalist
who has spent some time in the tropics expresses a decided
opinion to the effect that birds do not normally eat butterflies,
because he has never observed them doing so, it is incumbent on
us, before accepting his evidence as having any real scientific
value, to satisfy ourselves that he has made a systematic and
thorough investigation of the subject, and that his views are not
based merely on casual and inadequate observations. For in a
matter of this kind there is grave danger that absence of evidence
may be due simply to lack of observation. If a collector main-
tains that birds do not eat butterflies, we are justified in asking
him for @ full list (italics mine) of the other insects which he has
seen captured by birds. And I venture to think that a closer
inquiry of this kind would reveal the fact that most of the
negative evidence which has been brought up against the Select-
ionist interpretation of mimicry is really of little worth.”

Passing over the obvious reply that as it was the Selectionists
who first asserted that birds ate butterflies, it is their duty to
prove it if they wish their theory accepted, it would appear
that My. Marshall does not consider it necessary for an observer
to be very accurate as to the species captured before attributing
butterfly-catching propensities to certain birds.

In the ease of the Redstart we read that ‘ They take flies, gnats,
small butterflies, and all sorts of small two- and four-winged insects,
partly on the wing and partly at rest”; and again, “ It feeds on
flies, gnats, small butterflies, and various other kinds of small
coleopterous and other insects, caterpillars, etc.” Now if such
evidence is accepted, namely, that the Redstart eats butterflies,
without the necessity of naming the individual species captured
(though it might well be suggested that these small butterflies
were really small moths), it would appear only just that when
such an authority as Legge states that the food of the large Indian
Cuckoo-Shrike consists of “ cater pillars, grasshoppers and various
other kinds of coleopterous insects” without mention of butterflies,
that such should be regarded as sufticient evidence that butterflies
are not destroyed by it in sufficient numbers to cause any form of
mimicry. But, on the other hand, when we read of a bird feeding
‘‘on beetles and the many larger members of the insect kingdom
which affect Ceylon forests,” it 1s quite possible that such a one
would produce a struggle for existence among butterfiies. It is
only by adopting some such standard as this, faulty though it may
be, that we are likely to come to any conclusion. Asto the actual
observation of insects captured by birds, no one who has not

m2, LY.-COL. N. MANDERS ON 'THS

experienced it can form an idea of its difticulty. Let any one who
would appreciate it, watch a blackbird or thrush in his own garden,
and even with the best field-glasses he will be unable in the great
majority of cases to name the species of insect caught, or more fre-
quently than not the order to which it belongs. In the tropics the
difficulties are increased a hundredfold : fortunately in the case of
butterflies the task is lighter, and it is often easy to name the species
owing to the habit some birds have of holding the insect in the
bill for a few moments before swallowing it, and the lengthy
time it takes others either to tear off the wings after the manner
of Shrikes, or beat them off against the ground as is the custom of
the Robin family. Bee-eaters and Paradise Flycatchers nip off the
wings close to the body as neatly as if done by a pair of scissors.

A certain number of birds are migrants, such as the Cuckoo
and Swallow, which pass a portion of the year in Ceylon and the
remainder in northern latitudes; such birds, as pointed out by
Mr. Marshall, would have to learn the distastefulness or otherwise
of the butterflies that inhabit such dissimilar countries, and
their influence on the butterfly population of both areas would
have to be taken into consideration. These are, however, few in
number compared with the partial migrants, which move from
one part of the Oriental region to another; and the butterflies
there being of the same character, the lessons they have learnt in
one part of their distribution are valuable to them in another.
A great many of the resident insectivorous birds move in an
irregular manner from one part of the island to another according
as their food supply varies, and as this consists of insects alone it
is largely dependent on the rainfall. Such a movement cannot
be strictly termed a migration and may be a few miles only ; for
instance, the rainfall in Colombo is about eighty inches, twenty
miles off it is nearly two hundred; when there is a comparative
paucity of insect life at Colombo the birds, or rather some of them,
find plentiful sustenance by moving a few miles inland. Some
birds again, such as the Robins, remain in and about the houses
and gardens year after year, and others, such as the Green Bee-
eater, are never found in the wet portion of the island.

It is this slight to and fro movement which makes me sceptical
of any insectivorous bird in Ceylon, and probably in any tropical
island, being ever really hard up for food.

Mr. Marshall quotes Dr. Franz Doflein as writing: ‘‘ From the
observations which I made in the jungles of Ceylon, it is quite
incomprehensible to me how naturalists who have spent years and
tens of year’s in the tropics can deny the fact” [that butterflies
are frequently attacked by birds]. I had very little doubt when
I read this passage that Dr. Doflein was speaking of the north of
the island, and his recorded observations confirm this; I should,
however, be very surprised if he could say the same of the whole
island, more particularly the hill districts. One point to which I
wish more particularly to draw attention, is his suggested
immunity from attack of the magnificent Ornithoptera darsius,

VALIDITY OF SOME FORMS OF MIMICRY. 713

attention being drawn by him to “its slow, lazy and almost
unwieldy flight,” the characteristics of an unpalatable butterfly.
It does not occur or very rarely in the north of the island, where
a bird, the Paradise Flycatcher, is very abundant, but at Kandy,’
to which place every visitor goes, and where possibly Dr. Doflein
made his observation, the butterfly is very common and the bird
rare, though [ have seen it. But it so happens that at the foot of
the Nilgiris, both bird and butterfly inhabit the same district and
the former is an inveterate enemy of the latter, it eats numbers
of them by nipping off the wings and swallowing the body. The
slow, lazy flight may possibly advertise its unpalatability to some
birds, but it makes it the easier victim to the Paradise Flycatcher,
which in my opinion is the greatest enemy butterflies have in
this part of the world.

I have always experienced considerable difficulty in under-
standing how a distasteful butterfly has acquired a slow sailing
flight *. It is easy to see how swift flight could be brought about
by natural selection, but the converse is not so clear. Presumably
Eupleeas, Danaines, etc., have always been distasteful on account
of the poisonous or nauseous nature of the food-plants, and those of
slower flight, as in the case of the palatable kinds, would naturally
be first captured, and we can understand how a race of quickly-
flying evil-tasting butterflies would be evolved by natural selection.
But if quick flight is of assistance in enabling a tasty butterfly
to escape capture, I find it difficult to believe that a nasty one,
with equal powers of flight, would not be equally benefited. And
if this be so, slow flight for the purpose of advertising unsaleable
goods seems unnecessary and the method by which it has been
evolved very obscure.

When we come to study fast-flying butterflies in their native
haunts, we find in every case a different rate of flight according
to the hour and weather. If it is a cloudy morning they fly much
slower than they do in hot bright sunshine; the majority fly
their fastest after 10 a.m. till 3 p.m., often resting between 12
and 2. But in the early morning and late afternoon, these same
butterflies can often be captured with the greatest ease, as at this
time they are usually feeding. This is the case with the rapidly
flying Teracolus (fausta, danae, &c.) and many Papilios, and this
may account for the different opinion Dr. Doflein and I hold
regarding P. hector and P. polytes. He considers them to be of
swift flight, and so no doubt they are in the middle of the day,
and particularly when flying over bare ground from one patch of
cultivation to another; but on a dull day or early in the morning
or evening they are particularly easy to catch. If Dr. Doflein is
correct we have here an instance of a Miillerian combination of

* Mr. Marshall speaks of “the acquisition of unpalatability,” by which I presume
he means that a slightly nauseous butterfly has become more so by a process of
evolution, and this has been accompanied by an increasingly slower flight. There
is no proof, so far as I know, that a Huplea for instance has undergone any such
process, and the assuinption appears to be entirely theoretical, :

714 LT.-COL. N. MANDERS ON THE

unpalatable butterflies adopting or having a naturally quick
manner of flight which enables them more or less successfully to
avoid the tasting experiments of young birds. I mention this in
order to show the difficulties that beset us even in the field, and
how two observers may form a diametrically opposite opinion on
apparently such a simple matter as the flight of a buttertly.
Dr. Longstaff is no doubt correct in saying that P. polytes has a
quicker flight than P. hector.

This varying velocity of flight makes me somewhat sceptical
that swiftness has been evolved in order to escape capture, as an
enemy by selecting the opportune moment can effect the seizure
of a fast-flying butterfly as easily as one of slow and laborious
flight. ‘Those foes of butterflies, the Bee-eaters, feed during the
hot hours of the day, and Drongos in the morning, late afternoon,
and often after sunset.

Many butterflies are conspicuous under one set of conditions
and the reverse in another, even at the same time of the day.
Telchinia viole is said to belong to the most distasteful group
of butterflies. Its flight is slow and deliberate, and it is very
conspicuous when flying over a grass field; its bright brick-red
colour forms a strong contrast against the green, and it thus has
the characteristics of an inedible insect. In the blazing dazzling
sunshine on the dried-up plains of India its colour so matches the
soil that it is decidedly difficult to see, particularly the female
which is almost invisible. Its under surface matches admirably
the dried-up leaves of the bush on which it frequently takes up
its position for the night, and under these circumstances it has all
the characteristics of an edible insect. On a day in March this
butterfly was flying over the green gardens of Colombo, and three
days afterwards I met it at Trichinopoly, with a shade temperature
of 104°, dazzling sunshine, and scarcely a blade of grass to be seen.
It will probably be held by Selectionists that in certain cases
such as in extremely dry weather, even an inedible butterfly
requires concealment and that conspicuousness is beneficial to it
as advertising the worthlessness of its goods in the wet season,
when insects are abundant. I have given my reasons for
believing that in tropical islands there is no real scarcity of insect
life that cannot be made good by birds. It may also be argued
that its invisibility one moment and conspicuousness the next may
be of advantage to it, but if this be so, it is difficult to understand
why such an extremely distasteful insect as an Acrea is held to
be, should be obliged to pass through such a complicated process
of evolution.

In compiling the following list of birds, I have followed Oates
and Blanford, as their nomenclature is more modern than that of
Legge; where not otherwise indicated, the notes in inverted
commas are quotations from Legge.

Captain Legge spent eight years in Ceylon, and those who were
there with him, now few in number, have a clear recollection of
his knowledge, zeal, and painstaking industry.

VALIDITY OF SOMf FORMS OF MIMICRY. all

Ol

Fam. CRATEROPIDA.

Subfam. Crareropin®. (The Babblers.)

“ All feed on the ground like thrushes. They ... probably
derive no portion of their food directly from trees, the fruit they
occasionally eat being picked off the ground as they forage for
insects.” (Oates.)

1. The Southern Indian Babbler. Crateropus striatus.

“Its food is entirely insectivorous, and is mostly taken by
scratching among leaves and débris on the ground.”

2. The Ceylonese Babbler. C. rufescens. Indigenous.

“T found the stomachs of several examples killed in the month
of August to contain portions of a large black beetle which was
affecting the jungle in large numbers at the time.”

3. The Ashy-headed Babbler. C. cineretfrons. Indigenous.

“Delights in exploring the mossy recesses of fallen trunks, in
which humid spots it finds an abundance of caterpillars, bugs,
hemiptera, and coleopterous insects.”

4, The Ceylonese Scimitar Babbler. Pomatorhinus melanurus.
Indigenous.

‘¢Goes about in small companies searching for its insect food
on low branches or clinging w oodpecker fashion to the trunks or
large branches, about ne it jumps and twists itself with con-
siderable agility.” ;

Subfam. TIMELIIN a.

5. The Small White-throated Babbler. Dwmetia albigularis.
‘‘Tts food consists of the larvee of various insects and minute
coleoptera.”
6. The Ceylon Yellow-eyed Babbler. Pyctorhis nasalis.
Indigenous.
‘“‘T have cone found its food to consist of small coleoptera
and various minute insects.”
7. The Brown-capped Babbler. Pellerneum fuscicapillum.
Indigenous.

‘Tt feeds on the ground in dense thickets, picking up beetles
and insects from amongst decaying herbage : it rarely shows
itself in the open.”

8. The Black-fronted Babbler. Rhopocichla nigrifrons.

Indigenous.
“This modest but active bird frequents underwood, anata
and tangled jungle. .... subsisting entirely on various insects

and their larve.”

There is no indication among the Ceylon Babblers of any

716 LT.-COL. N. MANDERS ON THE

butterfly-eating propensity, but Mr. Frank Finn experimented with
an Indian species (Crateropus canorus), and came to the conclusion
that they distinguished in time between a tasty and distasteful
butterfly. The note I made at the time I studied his experiments
is as follows :—It is evident that they had no notion at first as to
what was palatable and what was unpalatable, but as the experi-
ments proceeded they learnt gradually to discriminate ..... I
conclude that as these birds with one exception were adult when
captured, they could not have undertaken tasting experiments
when young, otherwise they would have recognized a distasteful
species.

Subfam. BRACHYPTERYGIN ®.

9, The Indian Blue-Chat. Larvivora brunnea. A migrant.
«“ Appears to feed entirely on the ground.” (Oates.)

10. The Ceylon Arrenga. Arrenga blight. Indigenous.

“The food consists of various insects and in the stomach of my
specimen I detected the bones of a frog.” Mr. Oswin Wickwar
tells me that he found a species of snake, Aspidwra sp., quite four
inches long in the stomach of the bird shot by him.

11. The Ceylon Short-wing. EHlaphrornis palliseri. Indigenous.

“Found in thick brushwood feeding on the ground.” (Oates.)

“ Tt feeds on ants and other minute insects and to some extent
on small seeds.”

12. The Indian White-eye. Zosterops palpebrosa.

13. The Ceylon White-eye. Zosterops ceylonensis.
These species have the same habits as those in Bourbon and
Mauritius.

Subfam. LiorRicHIN&.

14. The Fairy Blue-bird. Jrena puella.
“Tt feeds principally on fruit.” (Oates.)

15. Jerdon’s Chloropsis. Chloropsis jerdont.
“ Seeds may often be found in its stomach, though they are not
so generally partaken of as insects.”

16. The Malabar Chloropsis. C. malabarica.
“‘ Lives on fruit and insects, chiefly the latter.”

17. The Common Tora. “githina tiphia.

‘“‘T have occasionally seen it dart out and seize a passing moth
or butterfly on the wing and alighting again swallow it whole, a
habit which is testified to by the large Mantide and other winged
insects which are often found in its smallstomach.” ‘ Frequents
orchards ..... feeding on insects which it finds among the
leaves.” (Octes.) ; i

VALIDITY OF SOME FORMS OF MIMICRY. Ae

Subfam. BRACHYPODIN2.

18. The Southern Indian Bulbul. Hypsipetes ganeesa.

“Tts diet consists of fruits, seeds and berries....... 1b,
however, also feeds on insects, and I have observed it occasionally
dart at them from its perch, although its usual manner of cap-
turing them is to seize them from the branches of trees, to which
it will sometimes cling after the manner of a Tree-creeper.”

19. The Madras Red-vented Buibul. d/olpastes heemorrhous.
‘Weeds mostly on fruit.” (Oates.)

20. The Yellow-browed Bulbul. Jole ictertca. Indigenous.

“T have found it to be more insectivorous than frugivorous.”

‘Tt wanders about in small flocks, feeding almost entirely on
fruits and seeds.” (Bourdillon.) ‘In all the specimens I have
examined I have found fruit only in its stomach, but from the
strong bristles at the base of the bill I suppose it may, at certain
seasons, partake of insects.” (Jerdon.)

21. The Black-capped Bulbul. Pycnonotus melanicterus.
Indigenous.
‘Tt is chiefly insectivorous, small seeds are sometimes devoured
by it, and I have found snails of some little size in its stomach.”

22. The Yellow-eared Bulbul. Kelaartia penicillata. In-
digenous.

23. The White-browed Bulbul. Pycnonotus luteolus.

‘Tt is both insectivorous and frugivorous, chiefly the latter, and
there is nothing to which it is more partial than the seeds or
berries of the latana plant.”

Bulbuls are very frequently kept as cage birds, and have perhaps
been more often experimented with than any other tropical bird.
Those I kept years ago ate any butterfly given them, and I think
it is now generally acknowledged that those species which are
known to be insectivorous also attack butterflies, but they show
no discrimination in eating them.

Fam. Dicrurip#. Drongos or King Crows.
Ali the Drongos are known to eat butterflies.

24. The Black Drongo. Dicrurus ater.

‘¢ The principal food consists of coleoptera, grasshoppers, winged
termites, of which it is very fond, and ticks, which latter it takes
from cattle. It has been known to devour small birds.”

25. The Indian Ashy Drongo. D. longicaudatus. Migratory,
Proc, Zoou, Soc.—1911]1 No, L. 50

718 LT.-COL. N. MANDERS ON THE

26. The White-vented Drongo. D. leucopygialis. Indigenous.
“Tt is entirely insectivorous, its diet consisting chiefly of
beetles, bugs, termites and such like.”

27. The Ceylon Black Drongo. Dissemuroides lophorhinus.

Indigenous.
“Damp forests and even their most gloomy recesses are fre-
quented by this fine bird...... It feeds on beetles and the many

larger members of the insect kingdom which affect Ceylon
forests.”

28. The Larger Racket-tailed Drongo. Dissemurus
paradiseus.

‘“‘ Feeds chiefly at dusk when the bats come out.”

The distribution of this family in the island is to be noted.
The first species is confined to the Jaffna peninsula and north-
west coast: I found it abundant on the opposite Indian coast in
March. The second is migratory and does not breed in the island.
The third is of general distribution. The fourth is confined to
the heavy forests of the Western Province. The fifth to the
North and Eastern Provinces.

Fam. Syuvirp#. Warblers.

29. The Indian Great Reed-Warbler. <Acrocephalus stentoreus.
“The diet consists of small flies and minute insects.”

30. Blyth’s Reed-Warbler. 4A. dwmetorwm. Migrant.

31. The Rufous Fantail. Cisticola cursitans.

“The diet of this species in Ceylon consists of many sorts of
small insects and caterpillars.” ‘‘The indigestible parts of the
food, which consists of small beetles, caterpillars and little snails,
are thrown up in pellets.” (Jerdon.)

32. Franklin’s Wren-Warbler. Franklinia gracilis.

“The food consists of small insects, which it picks up among the
dead wood to which it is so partial.”

33. The Broad-tailed Grass-Warbler. Schenicola platyura.
“‘ Feeds on the ground.” (Ouates.)

34. The Greenish Willow-Warbler. Acanthopneuste viridanus.
A migrant. ;

35. The Ashy Wren-Warbler. Prinia socialis.

“‘TIts food consists of insects; but occasionally I have found
small seeds in its stomach.”

36. The Jungle Wren-Warbler. Prinia sylvatica.
*Tts food consists of small coleoptera and other minute insects,”’

VALIDITY OF SOME FORMS OF MIMICRY. 719

37. The Southern Wren-Warbler. Prinia jerdon.
‘“‘Tt is purely insectivorous.”

Fam. Laniip&. Shrikes.
Subfam. LANIINA.

“These birds live entirely on insects, the Tree-Shrikes oc-
casionally seizing a small bird or mammal. Some descend to the
ground to seize their prey, a few catch insects entirely on the
wing, and others, again, search branches and leaves for their
food.” ( Oates.)

38. The Rufous-backed Shrike. Lantus erythronotus.

Sew ocally bicdeen ener feeds on grasshoppers, which it entraps
on the ground, and also preys on Mantide and dragonflies.”

It almost certainly eats butterflies. I haveseen it occasionally
in Colombo.

39. The Brown Shrike. Lanius cristatus. A migrant.

Certainly eats butterflies. See experiments on birds in
Colombo (p. 737).

40. The Black-backed Pied Shrike. Hemipus picatus.

“They are rather flycatchers than shrikes in their habits . .
. continually darting out and seizing insects on the wing.”

(Oates.)

41. The Common Wood-Shrike. TZephrodornis pondicerianus.

“Moths and small butterflies form a considerable portion of
its food.”

42. The Orange Minivet. Pericrocotus flammeus.

“ Its diet consists of small butterflies and various winged insects.
In the woods of the Horton Plains I saw it catching insects
in the moss with which the trees are entirely covered in that
damp region.”

Mr. Ormiston tells me that he has known a small flock of these
birds completely clear off some dozens of Papilio polytes larve,
which he was hoping to rear on the fruit trees in his garden.

43. The Black-headed Cuckoo-Shrike. Campophaga sykesi.
‘‘ Its favourite food is caterpillars and other soft insects.”

44. The Large Cuckoo-Shrike. Graucalus macii.

“‘Tts food consists of caterpillars, grasshoppers and various kinds
of coleopterous insects.” Hodgson states its food to be ‘‘ Mantidee
Scarabeei, berries, vetches and seeds.”

45. The Little Minivet. Pericrocotus peregrinus.

“Tt feeds upon various larve (its favourite food) and small
insects.”
50*

720 LT.-COL. N. MANDERS ON THE

Subfam. ARTAMIINA.

“They catch their food entirely on the wing, either by darting
on it from a fixed perch or by flying about after the fashion of
swallows.” (Oates.)

46. The Ashy Wood-Swallow. <Artamus fuscus.

This bird has been frequently quoted as having been seen by
Colonel Yerbury to catch several Huplea core.

Fam. SturNip#. The Mynahs.

“They feed chiefly on the ground on insects and worms, but
they are fond of fruit and berries, which they pick off trees.”
( Oates.)

47. The Black-headed Mynah. Temenuchus pagodarum.
Widely distributed and common.

48. The White-headed Mynah. Sturnornis senea.
A rare and local resident.

49, The Common Mynah. Acridotheres tristis.
See experiments on birds in Colombo (infra, p, 740).

Fam. Muscicapip#. The Flycatchers.

‘The Flycatchers feed on insects which they either catch on the
wing, starting from a perch to which they usually return several
times, or by running with the aid of their wings along the limbs
of trees. They seldom or never descend to the ground.” (Oates.)

50. The Indian Red-breasted Flycatcher. Stphia hyperythra.
Partial migrant.
See experiments on birds at Nuwara Eliya (infra, p. 735).

50 (a). Tickell’s Blue Flycatcher. Cyornis tickelli. Food ?

51. The Blue-throated Flycatcher. Cyornis rubeculoides.
Partial migrant.

52. The Ceylonese Dusky-blue Flycatcher. Stoparola sordida.
Indigenous.
See experiments on birds at Nuwara Eliya (p. 735).

53. The Brown Flycatcher. Alseonaa latirostris, Migrant.
The habits of this bird are well known.

54. Layard’s Flycatcher. <Alseonax muttwi. Partial migrant.

‘‘ In the stomach of one example I found much larger insects
(moderately sized coleoptera) than I expected to find captured by
so small a bird,”

VALIDITY OF SOME FORMS OF MIMICRY. 72)

55. The Grey-headed Flycatcher. Culicicapa ceylonensis.
Jerdon says it feeds on small insects.

56. The Indian Paradise Flycatcher. TZerpsiphone paradist.
Partial migrant.
See correspondence (infra, pp. 728 & 730).

57. The Indian Black-naped Flycatcher. Hypothymis azurea.

Mr. Oswin Wickwar has given me the following interesting
note :—

‘“‘T watched two flycatchers of this species diving into a small
pond evidently in search of some aquatic insect. They both dived
in the most determined manner about five or six times, and,
although I looked carefully, I could not find any insects on the
surface of the water. They did not swoop down and just touch
the surface of the water in the manner of swallows, but deliberately
dived in with a splash like a kingfisher ; there was a momentary
pause, and then they fluttered back to the same perch or one near
about where they started from.”

58. The White-browed Fantail Flycatcher. Rhipidura
albifrontata.
“The chief food consists of mosquitoes and other small dipterous
insects, as also the small Cicadella.” (Jerdon.)

Fam. TURDID&.
Subfam. Saxtcotinz. The Chats.

‘“‘The Chats feed entirely on insects they capture generally on
the ground from a fixed perch, such as the summit of a stone, a
stalk of grass, or a branch of a bush, and then return at once to
their post of observation.” (Oates.)

59. The Southern Pied Bush-Chat. Pratincola atrata.

“The food consists of insects and larve of various kinds, which
they take chiefly on the ground.” This bird is also known as the
Nuwara Eliya Robin; it has very much the habits of the
Stone-Chat.

60. The Black-backed Indian Robin. Thamnobia fulicata.

Has been known to capture Veptis varmona; it usually feeds
just at sunset and as long as there is light.

61. The Magpie Robin. Copsychus saularis.
See experiments on birds in Colombo (p. 737).

62. The Shama. Cittocincla macrura.

“Those shot in Ceylon seemed to be entirely insectivorous,
the food consisting of small beetles, ants, flies, etc.”

722 LY.-COL. N. MANDERS ON THE

Subfam. TurDIN«».

63. The Ceylon Blackbird. Merula kinnisi. Indigenous.
Has almost precisely the habits of the English Blackbird.

64. The Ceylon Thrush. Oreocincla imbricata. Indigenous.

“Decidedly an uncommon bird.... it appears to feed on
insects which it procures beneath fallen leaves.” “‘ Thwaites says
it scratches much in rubbish thrown out at the border of his
plantation.”

65. “The Spotted-wing Thrush.” 0. spiloptera.
““Generally to be found in thick damp jungle picking up pups,
coleoptera and other insects.”

Three species of Swallow occur; one, the Common Swallow
(H. rustica),is migratory, the other two have similar habits.

Fam. MovTAciILuip#. Wagtails.

There are four species of Wagtails, three of which are migrants ;
the fourth, the Large Pied Wagtail, has the same habits as the
rest of the family.

66. The Indian Pipit. Anthus rufulus.

“Weeds on worms and various terrestrial insects and likewise
partakes of small grass seeds.” I have noticed it feeding on green
Aphides, and once saw it capture a Lycznid, a species of Zizera.

Fam. CorAcitAap#. The Rollers.

67. The Indian Roller. Coracias indica.

‘“‘T have on several occasions seen one pursue an insect in the
air for some distance and when the winged termites issue from
their nests after rain, the Roller, like almost every other bird,
catches them on the wing.” (Jerdon.)

“Its food is chiefly large insects, grasshoppers, crickets,
Mantids, and even beetles, occasionally a small mouse or shrew.”

(Jerdon.)
Family MEROPIDS.

68. The Common Indian Bee-eater. Jferops viridis.

69. The Blue-tailed Bee-eater. IL. philippinus.

‘A winter visitant..... it feeds on wasps, bees, dragonflies,
beetles, and even butterflies.” (Oates.)

70. The Chestnut-headed Bee-eater. Jf. swinhoei.

“ Locally distributed throughout Ceylon.” Not found in the
hill country.

These three species are well-known as butterfly-eaters; see
correspondence (p. 727).

VALIDITY OF SOME FORMS OF MIMICRY. 723

Fam. ALCEDINIDS#. Kingfishers.

71. The White-breasted Kingfisher. Halcyon smyrnensis.

“Tt occasionally, but rarely, catches fish by plunging after them,
it lives chiefly on insects and small lizards and sometimes on mice
and land crabs.” (Qates.) ‘‘ It subsists on lizards, grasshoppers,
locusts, and even small snakes.”

Fam. CypseLip#. Swifts.
72. The Alpine Swift. Cypselus melba.

73. The Common Indian Swift. C. affinus.
These birds have all the habits of the common European Swift. _

74. The Palm Swift. Zachornis batassiensis.
‘“‘ Feeds chiefly at dusk.”

75. The Brown-necked Spine-tail. Chetura indica.

“Mr. Carter says that those he shot had fed on beetles, green
bugs, sand-wasps and grasshoppers.” Mr. Spurway informs me
that he has more than once seen it snap up a butterfly. Legge
says it is fond of termites.

76. The Indian Crested Swift. J/acropteryx coronata.
I know nothing about the feeding habits of this bird.

Fam. Trogonip#&. The Trogons.

77. The Malabar Trogon. Harpactus fasciatus.

Mr. Butler has seen a large moth brought to the nest he was
observing.

“ Tt feeds chiefly on beetles, moths or cicades ; but it occasionally
feeds on insects on the ground.” (Blanford.)

Fam. Cucuuip#. The Cuckoos.
‘They feed chiefly on caterpillars and soft insects.” (Blanford.)
78. The Cuckoo. CO. canorus Migrant.
79, The Small Cuckoo. (C. poliocephalus. Migrant.

80. The Indian Cuckoo. C. mucropterus.
‘¢ Feeds on caterpillars.”

81. The Indian Plaintive Cuckoo. Cacomantis passerinus. A
migrant.
«“ Tt feeds on caterpillars, coleoptera and other large insects, and
may often be seen taking them on the ground.”

M24: LT.-COL. N. MANDERS ON THE

82. The Banded Bay Cuckoo. Penthoceryx sonneratis.
‘Feeds on coleoptera, Mantide, and caterpillars.”

83. The Drongo Cuckoo. Surniculus lugubris.

‘Locally dispersed .... the diet is mixed, consisting chiefly
of caterpillars and beetles but often combined with various seeds.”

84. The Pied Crested Cuckoo. Coccystes jacobinus. Migrant.

85. The Red-winged Crested Cuckoo. C. coromandus. Migrant.

‘‘The stomachs of those I have procured contained beetles,
grasshoppers, Mantide, and other large insects.”

Fam. FALCoNnIDs#. The Falcons.

86. The Kestrel. Zinawneulus alaudarius.

Has the same habits as the English bird, which has re-
cently been shown in some instances to have butterfly-eating.
propensities.

The above birds may be tabulated as follows; the arrangement
is artificial, and no doubt there are errors, but it is convenient.
Some may be inclined to transfer the Flycatchers from group 3
to group | on the evidence brought forward by Mr. Marshall in
the case of the European Flycatcher. I have put them in the
third group, as close observation for two months showed me
that two species at any rate only eat butterflies under unusual
circumstances.

May take Butterflies, |

ST ihre ta ee gee e afer atare |
| The Common Tora 1 | Chats and Robins... 4] Flycatchers ......... 8
| Dron Ose. ye eee eee | Swallows ............ 3) Sbabblersipaes sees
SIRAUEE” eceaneehbcouns eGapncencccuts Gy WES ee sch encodes ZI) WAVMIDERS. “lecodheoc: 9 |
The Orange Minivet ............ 1 | SURORQOIN soocccceaccdues Jb || ISGUNIS conpsadéeseonovace 2
The Ashy Wood-Swallow ...... 1 | Cuckoos oueeeeesee., 8} The Little Minivet 1
The Paradise Flycatcher _...... il DAMM socscune 9052 6 | ChloropSis ............ 2)
Way) OW Pecans ndoastcasteopcanccsca:’ L The Kestrel ......... 1
Beezeaters.... canoes oe eee: | Miynahs” jo. ase a
The White-breasted Kingfisher 1 | The Indian Pipit ... 1
The Brown-necked Spine-tail ... 1 |
Toran... 21 | Toran... 31 ToTat... 30
4 of the above are either mi- 9 of the above are 6 of the above are |
grants (2) or partial migrants (2). | either migrants (4) or | either migrants (4) or
partial migrants (5). partial migrants (2).

I would suggest this as a convenient place for a perusal of

VALIDITY OF SOME FORMS OF MIMICRY. 725

Mr. Marshall’s paper above quoted, more particularly that part
devoted to the Indo-Malayan region.

For the sake of convenience I shall designate the birds in
Group 1, “The Butterfly-Eaters,” and the evidence in support
of this term is given below under each species.

1. The Common Tora and Butterflies.

This bird belongs to the Bulbul family, another name for it
being “ The Common Bush Bulbul.”

I have frequently watched this bird and can only confirm
Legge’s observations. Being so closely allied to the Bulbuls it
is probable that, like its near relations, it is an indiscriminate
feeder.

2. King Crows (Drongos) and Butterflies.

“On the 14th instant I was with Mr. C. B. Antram in a very
small patch of wood surrounded by grass downs within a few
miles of Ootacamund. Running through this wood was a foot-
path, and this path was in one place a few feet long thickly
strewn with the wings of butterflies; on either side of this, for
some yards along the path, were scattered wings. Just above
this place three Drongos (Dicrurws) were to be seen on the trees.
The weather during my visit was misty, cold and rainy, and hardly
any butterflies were on the wing ; consequently I had not the
opportunity of seeing the birds hawking them. Dragonflies were
abundant and the Drongos made frequent excursions after them,
all unsuccessful so far as I saw. On examination, the wings were
found to belong to the following species:—Danais limmiace (or
septentrionis, or both), Danais aglea, Danais chrysippus, Huplea
core, Huplea coreta, Hypolimnas bolina 2 , and Catopsilia crocale.
At least 90 per cent. of the wings belonged to D. limniace or
septentrionis. There were several wings of Huplea, amongst
which I found those of the males of both core and coreta. The
remaining species were each represented by only one or two
wings. These Danaine are common in those parts, and in
sunny weather would be passing in numbers through and over the
wood, and the most common would almost certainly be limniace,
septentrionis, core and coreta. D. chrysippus would be commoner
probably on the outskirts of the wood. The only other butterflies
about in any numbers when I was there, were Argynnis castetsi
and Colas nilgiriensis, both eminently insects frequenting the
open country, and they would seldom or never be found in any-
thing heavier than scrub jungle. Some of the wings were
obviously quite fresh, others had been beaten on the ground by rain.
The three Drongos were almost the only birds about, and no
other birds that we saw were likely to be capable of capturing
these butterflies. It appears to me that the Drongos were
certainly the cause of this extensive shower of wings; the only
other possibilities are lizards or mantids, but no lizard, I should

726 LY. COL. N. MANDERS ON THE

imagine, has either the activity or voracity to make away with
sucha great number of butterflies, even assuming that they settled
within its reach. A mantid is even less probable, and I much
doubt if there is one large enough to tackle Danainz or Hypo-
Limnas in the Ootacamund region.

“One would not be surprised to find an occasional Danas or
Euplwa sampled by a bird, but to see evidence of a systematic
onslaught on butterflies which are so universally looked upon as
leaders of the army of distasteful insects, and which are so widely
mimicked by numerous ‘unprotected’ butterflies and moths,
tends to make one sceptical of the accepted theories founded on
the alleged value of this distastefulness. It is true that they may
disagree with other birds, lizards, &c., but if one enemy alone can
effect such wholesale destruction upon them, their immunity from
death by violence is so seriously impaired that it seems to me that
their numerous imitators amongst the ‘ Swallowtails,’ &c., are
simply asking for trouble.

“There was no lack of other food for the Drongos, and it can
only be assumed that they found the Danaine very much to
their taste. One can hardly think of a morsel more apparently
unpalatable than Telchinia viole, yet I saw a Bulbul feed its
young with one within a few feet of me. It would be interesting
if entomologists would give any evidence in the matter which
they can. I have seen the wings of Hypolimnas misippus some-
times scattered on the road near trees in considerable numbers,
and on two occasions I have seen the wings of Charaxes imna ;
this I imagine was not caught on the wing; if so, I must con-
gratulate the bird on its power of flight.

H. Lestre ANDREWES.” —
Barwood Estate, Nilgiris, 20th October, 1910.

(Journal Bomb. Nat. Hist. Soc., vol. xx. p. 850.)

This interesting observation may be held to support Mr.
Moulton’s proposition (Trans. Ent. Soc. Lond. 1908) that the
Kupleas of S. India are a Miillerian combination formed for
mutual protection against the onslaught of insectivorous birds ;
but evidence is required that at one time they differed materially
from their present day appearance.

I took the few notes following at Coonoor, 6000 feet, Nilgiris,
S. India, in April 1910. I asked my collector, a half-caste who
had spent all his life in the jungles, whether he had ever seen a
bird catch a butterfly, and he immediately said he had, describing
the Paradise Flycatcher very accurately. He said they eat the
brown butterflies (1. core and coretw) and white ones. While he
was speaking I happened to open an envelope containing Danais
septentrionis, and he exclaimed, ‘‘ I have seen them eat those too.”
He added the bird nips off the wings and swallows the body ;
also that they catch and eat the Ornithoptera. A few days after-
wards I was at Kullar at the foot of the hills, about thirty miles
from Ootacamund, where these birds and Drongos are common.
I saw a Drongo in a lane, and close together on the ground

VALIDITY OF SOME FORMS OF MIMICRY. (OAT

I found the wings of Papilio hector, Huplea core, and Danais
septentrionis, and a little distance further on another D. septen-
trionis. ‘These were the commonest butterflies about at the time.

12.4.10. Kullar, 1200 feet. Saw a Bulbul dart out at a
Papilio hector and miss it. Saw the following with notched wings :
P. demoleus, one or two Catopsilias, one or two Teracolus fausta,
several P. hector, many Junonia lemonias, these last usually with
oval pieces out of the secondaries, probably by Calotes.

10.4.10. Coonoor, Walking along a road I saw what I thought,
at first, was a leaf falling from a tree about twenty feet up, but on
going to look at it I found it was the fore wing of H. masippus 9,
diocippus form. It is curious that I have only seen one other
female during the many times I have been along this road. I could
not see the bird, but Bulbuls are plentiful. Found also on the
road the fore wing of Papilio sarpedon.

21.4.10. Kullar. Watched a Racquet-tailed Drongo for some
time hawking after flies. I did not see it chase a butterfly
though there were numbers ftying about, chiefly Euploeas.

Mr. T. N. Hearsy, Indian Forest Service, writes :—‘‘ Coimbatore,
GiOalONs. ks ee I have frequently seen the common green Bee-eater
(Merops viridis)and the King Crow (Buchanga atra) take butterflies
on the wing, the butterflies being Catopsilia pyranthe, C. florella,
Terias hecabe, and Papilio demoleus. The Bee-eater I have also
seen taking Danais chrysippus and Danais septentrionis, and I
remember to have been struck with their taste for those latter. .
I have also seen the Tree-Swift (Dendrochelidon coronata) take
Catopsilia pyranthe.” -

In another letter he mentions having seen Drongos attacking
dragonflies.

Mr. Ormiston, of Kalupahani, Hadumulle, Ceylon, 4500 feet,
writes :—“Kalupahani, 4.2.09. Of course I will try and get you any
notes I can on birds eating butterflies ....I can give you very
little assistance at present, as the only bird I have watched is the
Fork-tailed Drongo, who eats the whites during a flight and attacks
Kallima.

‘“‘ The Magpie Robin and black-and-white Flycatcher catch a great
lot of moths, but I have no notes of their taking butterflies. At
Ohiya bungalow, after a moth night, we used to bottle all the moths
we wanted, and then loose a tame Mynah who made short work
of the rest, but I never tried him with butterflies. I will try the
Robins in my garden, but the fact that they eat dead butterflies
will not prove that they catch them. Personally I do not think
birds make any appreciable difference to the number of butterflies
except possibly by eating them in the larval stage. It is quite
unusual to see a butterfly caught.”

“ Kalupahani, 3.1.10. I have kept my eyes open for butterfly-
eating birds, but I am sure that the slaughter, if any, must take
place in the larval stage. I have seen the Fork-tail Drongo
feeding on the flight whites and Kallimas, but that is all.

““T have seen a dragonfly catch and kill Zesius chrysomallus
and another fly killing Papilios, Lycenide, Syntomiide, with

728 LT.-COL. N. MANDERS ON THE

apparently preference for the last named. Some black-and-white
Flycatchers come close when I am beating for ‘plumes’ on the
road here, and catch lots of common Pyralids I put up. The
common Magpie Robin comes most mornings for the moths which
I have slain at the lamp and discarded, and I have seen a Sparrow
catch a gold-tail moth. But since you asked me a year ago
to look out, I have not seen half a dozen cases of birds eating
butterflies.”

“ Kalupahani, 26.6.10. I have kept my eyes open for birds eating
butterflies but have no new notes. During the last flight the
Fork-tailed Drongos were as usual feeding on the white butterflies,
but that is the only instance I have seen. Certainly few, if any,
birds eat the Terias family. I have watched them flying slowly
with Swallows, Drongos, and Flycatchers close to them and leaving
them alone. I fancy Bee-eaters take butterflies, but they are
searce here, and I have no data therefore to goon......

“ Kallima philarchus appears at Hadumulle in large numbers
and the flight lasts for about a week. The biggest flights are
’ very nearly always from about Christmas to the New Year, but a
smaller flight appears in July. About once in four years they are
especially common . . . They seem especially to favour Loquat trees,
but come readily to a mixture of jaggery (native sugar), beer and
rum. Instead of settling on the branches or trunk with their
heads towards the top, they seem usually to do the opposite, and
are therefore apparently very conspicuous; but the birds do not
seem to notice them till they move ren they come to sugar
they settle anyhow).

“The spot where I mainly catch them is about two acres of
Grevilleas planted with a few Loquats. When the flight is on,
the Grevilleas seem full of Forked-tailed Drongos, and as soon as a
butterfly moves a Drongo darts for him, but usually only takes a
big piece out of one wing. I have never seen the wings lying on
the ground, so fancy if the Drongo gets him he eats wings and all.
I have never, however, seen him catch one.”

“10.10.09. Paducka. Watched Drongos (Dicrurws leucopygialis)
hawking for flies ; though Mycalesis ceylonica and smali Lycznids
were flying plentifully beneath the birds, they did not take them.

“19.12.09. Paducka. Watched several Drongos and a Paradise
Flycatcher ; the former frequently caught small flies in the air and
occasionally came to the ground after bigger insects, but only once
did one catch a lepidopterous insect and this seemed to mea moth.
The Flycatcher took short flights on the ground picking up flies,
but certainly not a butterfly. Cameacross five fully fledged Ashy-
headed Babblers sitting all together on a branch ; they flew off only
when I approached quite close to them, with great chattering, very
much like the ordinary Babblers ; the old birds were hunting for
food in the thick bamboo jungle. This is very late in the year
for young birds.”

I give these merely as samples of negative notes; there is no
object in giving more.

VALIDITY OF SOME FORMS OF MIMICRY. 729

Mr. KE. Ernest Green writes :—
“* Peradeniya, 16th July, 1910.

“With regard to the capture of butterflies by birds, I was told
(in May last) by a lady who was staying with us, that she had
been watching the Drongos in these Gardens busily catching
butterflies. From her description, the victims seemed to be
species of Huplwa. She said that the birds bit off the wings, and
that the road was covered with the dismembered wings. I asked
her to collect some of the wings for identification, But, in the
meantime, they had either been swept or blown away, and
she could only produce one or two wings of Papilio jason and
Jamides bochus.”

The common brown Eupleea can hardly be mistaken for any
other Ceylon butterfly.

3. Shrikes and Butterflies.

Dr. Willey, F.R.S., writes:—‘‘ The late Grant Allen stated
positively that among the animals which he had seen in Butcher-
bird’s larders were mice, shrews, lizards, robins, tomtits, and
sparrows ; but he added that in spite of its occasional carnivorous
tastes, the Shrike is at heart an insect-eater.”

The few experiments I have been able to make leave little
doubt in my mind that they make little or no selection in their
butterfly diet. See below (p. 737).

4. The Orange Minivet and Butterflies.

I know nothing about this bird’s provender other than already
given ; Mr. Ormiston’s observation is, however, suggestive. The
larva of Papilio polytes lives in its earlier stages exposed on the
upper surface of the leaf of the orange or citron, looking exactly
like a bird’s dropping; when irritated it shoots forward two fleshy
“horns” emitting a pungent smell of orange, which is highly
disagreeable. I presume this is derived from the food-plant, and
if this be so the larva in all probability has a taste of unripe orange,
and consequently it would seem that the bird’s palate is not highly
educated and its taste in butterflies probably not selective.

5. The Ashy Wood-Swallow and Butterflies.

Mr. Walter A. Cave writes :—
“Colombo, 21st October, 1910.

‘“‘T am sorry I cannot help you much in regard to the question
of butterflies being taken by birds. On one occasion I observed
an Ashy Wood-Swallow (Artamus fuscus) tearing the wings off a
butterfly, then swallowing the body. This was in Peradeniya
Gardens a year or so ago. ‘There were many of these birds, which
are allied to the Shrikes, hawking over the Maha Weliganga river.
f did not see this particular bird actually catch the butterfly, but
I have every reason to believe it did, because I had a good pair

730 LT.-COL. N. MANDERS ON THE

of prism binoculars focussed on the bird as it alighted in a tree.
As I have said, the wings were first stripped, and as they fell I
was able to make them out. I am not well acquainted with
butterflies, but to make my notes complete I referred to the
Museum and determined the butterfly as the commonest brown
one we see about the roads here, and I put it down as Huplea
core,

“On another occasion I saw a Common Swallow (Hirundo
rustica) take a small white butterfly, but I was too far away, and
without my glasses, to determine the species. The Swallow was
hawking near a piece of very dark jungle, near the Nuwara
Eliya lake, and the white butterfly was very conspicuous against
the background.”

Colonel Yerbury’s note of this bird being seemingly partial to
Euplea has already been alluded to.

6. The Paradise Flycatchers and Butterflies.

Mr. John Pole, Scarborough, Maskeliya, 6000 feet, writes :
“ Maskeliya, 13.3.09. . . | seem to recall the attacks of the Odonata
and even Asylus (Diptera) on butterflies, but I never remember an
instance of a bird attacking one—I have watched the little Tailor-
birds eating the larve of Veritas hecube and that with seeming
distaste, and the Flyeatchers at work on Diptera from the shelter
of some leafy tree; I have had so large an insect as Phyllades
consolisma taken from under my very nose by a Drongo, have
had moths beaten from a fence in the daytime stolen by Swallows
ere I could net them, but have never seen a bird in Ceylon carry
off a butterfly. Im England I have seen a Swallow carry off
Papilio machaon whilst 1 was following it...I came out to this
Island in 1871.”

‘“‘Maskeliya, 17.3.09. Since writing on 13th we have had for two
days (16th and 17th) flights of butterflies, the first I have seen
this year. There has beenin my garden for the last three months
a bird, which generally goes by the name of the ‘Cotton Thief’
(the Paradise Flycatcher). This bird occupies a jak tree within
twenty feet of my window, and for the last two days he has been
obtaining all his meals from the flights of butterflies, and although
I have never actually seen him catch one, I have seen him
circling from the tree in pursuit and the wings of the insects he
captures falling around the base of the tree within a radius of
twenty feet. Should they be of any service to you, I can send
you the wings of Appias paulina and varieties ¢ and @? as
follows :—

Upper left wing, 4 g 2 @.
Upper right wing, 5 d 59.
Lower left wing, 5 Q.
Lower right wing, 2 3.

‘“« Maskeliya, 14.4... . I have only one more species of butterfly
to give you as its food— Papilio agamemnon, and this is the only

VALIDITY OF SOME FORMS OF MIMICRY, 731

butterfly to my best belief the bird has taken since my previous
letter on the subject. The bird is rare here ....We have had
no flights of the yellow and white Catopsilias this year—so no
wings ene the Appias victims were mostly the white forms 3g
and 92.”

7. The Roller and Butterflies.

Tam unable to add much regarding this bird’s dietary. It is
found only in the dry northern districts and is uncommon. In the
plains of India it is a familiar occupant of the telegraph-wire, and
I have often seen it chasing and no doubt catching various insects
from such a perch. I ‘have no doubt it catches butterflies, and
more than once am sure I have seen it do so, though I am unable
to name the species captured. Judging by its very varied menu,
I have little doubt that it pays little heed to the species it
manages to catch, but of this I have no direct evidence.

8. Lee-eaters and Butterflies,

Mr. Fred. Lewis, a well-known ornithologist, writes:—-‘‘ Colombo,
Sula ODN Ausra. 1 have noticed Swinhoe’s Bee-eater in particular
hawking after the common so-called ‘ Adain’s Peak butterfly ’
[ Catopsilia and App-as|, and it appears to prefer the white one to
the larger yellow fly. It does not, so far as I am aware, take any
of the large brown butterflies often to be found with the above
named, I have watched the bird when quite a selection of flies
could be made, but beyond taking the white and an occasional
yellow, I have never seen it feed on others. I am not prepared to
say, however, that the Bee-eater does not eat any other butterflies
than the two mentioned.

‘“¢ Our common black King-Crow appears to select the same flies,
taking them on the wing in the same way as the Bee-eater.
Swallows do not, so far as my observations go, ever touch any
butterflies.

“Tt isremarkable, however, on such occasions as one finds in the
dry zone, when vast masses of these ‘Peak’ whites and yellows
congregate round some wet pool or damp ground, that King-Crows
are not found taking the opportunity of a‘ square feed.’ I infer
therefrom that the butterfly is only a ‘side dish’ and not a
regular item of food. . . 1 cannot recall an instance of young birds
being fed with butterflies. I suspect the difficulty of swallowing
the wings is the reason, for I cannot say I have ever found small
nestlings with anything so difficult of mastication in their
mest,”

In another letter Mr. Cave writes :—

“ Belvedere, Colombo,
6th March, 1911.

“In reply to your letter I am sorry to say I have nothing
further to report on the subject of the capturing of butterflies by

732 LT.-COL. N. MANDERS ON THE

birds. I have the subject always in mind when I happen to get
out, and my friend Mr. Symons, of the Government Training
College, is also on the look-out, andif we should notice anything
it will be reported to you.

‘“‘ Being keen on birds there is very little that escapes my
notice when I happen to be out, and I must say the subject on
which you write is very exceptional so far as my observations go.
At Christmas time I motored round the south coast to Galle,
Hambantota and from thence to Haputale. The butterflies were
there in myriads, nearly all the way—none of us had ever seen
anything so remarkable in our lives. There were literally clouds
of butterflies—in fact we remarked that we now knew where all
the butterflies came from which used to appear here on migration
in the N.E. monsoon. But neither Mr. Symons nor I saw a
bird attempt to catch a butterfly, and we saw a large variety of
birds including Bee-eaters, Swallows, and Swifts.”

An argument of some force against the frequency of butterfly
victims may be advanced by the different behaviour of birds in
the presence of a flight of locusts and a migratory flight of
butterflies. I have been fortunate enough to witness both, and
the difference is very striking. In the former, every kite, crow,
and insectivorous bird in the district follows the locusts, gorging
to repletion; and it is a very remarkable sight the numbers, I
might almost say flocks, of birds following the swarm. In a
migratory flight of butterflies, on the contrary, with the exception
of a few Bee-eaters and Drongos, birds are conspicuous by their
absence.

Mr. Oswin Wickwar, F.E.S., sends me the following note :—
‘“When shooting in the Northern Province in May last, I was
crossing the dry bed of a river when a Bee-eater (Merops viridis)
swooped down and caught a butterfly on the wing within a
couple of yards of my face. The instant it snapped it up, the fore
and hind wings of the butterfly floated down in front of me, and
enabled me to prove its identity, Papilio polytes 3. This was
about 11 a.m. The following morning about 7 a.m. I had occa-
sion to cross the same spot, and saw what was probably the same
bird perched on a twig; I had hardly seen it, when it flew down
and entered a hole in the bank, but came out again in a few
seconds. This was evidently its nest, and I was anxious to look
for remains of insects, nor was I disappointed. By introducing a
twig, the hole seemed to be about two feet deep and to travel
in a horizontal direction, so a start was made to cut away the
bank, and before going a foot three insects were unearthed,—a
* humming-bird moth (MJacroglossa sp.?) which was still alive, a
‘skipper’ (Swastus grenvius) also alive, and a small green beetle
with white spots on the elytra. All these were secured and taken
away. The hole now took a turn and went a downward course
for about two feet, which meant cutting away a huge piece of the
bank to get at the nest, so I had to leave it,”

VALIDITY OF SOME FORMS OF MIMICRY. Gas

The following extract is of much interest ; it is from the ‘ Spolia
Zeylanica,’ 1910 :—

<< Bee-eaters as Fish-eaters.

“There are a pair of Chestnut-headed Bee-eaters (JJerops
swinhoet), which nest pretty regularly in a steep bank on a road
below my bungalow, and about 150 yards distant from my pond.
Almost any bright afternoon, between 2 and 3 p.m., they may be
seen fishing in the pond. They come down from a dead tree,
which stands on a knoll some 50 yards away ; sometimes hovering
for a moment over the water to locate their prey, but more
commonly marking it in their swoop, and dashing headlong into
the water like a kingfisher and very rarely missing their fish. IL
have seen the pair account for a dozen fish in as many minutes;
all quite small fry.

“When there is a flight of white butterflies on, these birds
devote most of their attention to them throughout the day, but
on warm bright days nearly always have a go at fishing in the
afternoon.

“‘T have always hitherto associated Bee-eaters with the one diet
of insects ; and I could not quite trust the accuracy of my eyesight
until I brought a strong pair of field-glasses to bear on the actors
at the short range of 15 to 20 yards. I think it probable that
many so-called insectivorous birds change their diet when some
chance has put them up to the taste of a new article which
happens to suit them. . .

‘¢ Since the pair, which I convicted of fishing, hatched out their
young, they have abandoned their fishing expeditions and may be
seen sitting on the tree facing their burrows catching insects
(chiefly white butterflies) to feed their nestlings.

E. Gorpon ReEgvss.”
Wiltshire, Matale, May 7, 1910.

9. The White-breasted Kingfisher and Butterflies.

1 have often seen this bird eating grasshoppers, and on one
occasion a butterfly which I was too far off to identify. Small
frogs and lizards, some of the latter of quite a respectable size, I
have also seen taken. Dr. Willey writes, ‘I remember being
much surprised, many years ago, to find a Kingfisher’s stomach
full of insect remains.” With such a mixed dietary its taste
for butterflies is probably impartial.

Sparrows and Butterflies.

Mr. R. D. Hodgins writes :—‘“ April 1911. These birds have
built their nests about my bungalow here at Matale, so I have
plenty of opportunity of watching them. JI have on three

Proc. Zoou. Soc.—1911, No. LI. 51

734 LT.-COL. N. MANDERS ON THE

occasions noticed the birds hawk and catch butterflies in mid-air,
and in two cases the butterflies caught were taken off the tree
tops, but I could find no trace of them on the ground.

‘“‘The flies on one of these occasions were brown and on the
other white, very like the common white cabbage butterfly of
England, but I was some distance away and couldn’t see them
properly [probably Huplea, and Catopsilia or Appias].

““On the third time the butterfly was caught while passing the
bungalow one sunny afternoon in February, but appeared to be
released the moment later and dropped to the ground. Only its
abdomen was taken and this was nipped off neatly at the waist.
These wings I collected and herewith enclose | Papilio sarpedon :
the butterfly was otherwise in remarkably good condition ; it is
a very rapid flier. |

‘“‘T have often seen a sparrow dive and catch a feather floating
in the air in a similar way to that in which a swallow does.
Whether the feather is mistaken for a butterfly or vice versa I
don’t know.

“In the case of Ceylon native birds, I have been unable to
detect any catches of butterflies, but have noticed that two of the
species will take and devour moths...... This bird [the
White-bellied Drongo| I have often seen catching flies from the
branch of a tree or telegraph-wire with downward swoop on to
the victim. I have often seen it catch small moths up to about
one inch in length, which it seemed to devour, wings and all, after
returning to its perch.”

It would be as well to refer to the distribution of the “‘ Butter-
fly-eaters.” The Drongos have already been alluded to; only
one is of general distribution. The Common Iora is widely
distributed. The Shrikes with two exceptions are found in the
wooded country of the upper and lower hill districts, the other
two I have seen only in the low country.

The Ashy Swallow-Shrike is a partial migrant in the island ; it
appears in small flocks in Colombo during the north-east monsoon,
at other times it is more an inhabitant of the north. The Roller
is found only in the dry districts, so also are the Bee-eaters,
particularly the Green Bee-eater, which is never found in the wet
country or above 300 feet. Swinhoe’s Bee-eater occasionally —
ascends higher, and the migrant Philippine Bee-eater appears for
a short time on the wet western coast at the break of the north-
east monsoon, but soon retires to the dry northern districts. The
White-breasted Kingfisher and the Brown-necked Spine-tail are
of general occurrence.

Tf this distribution is carefully studied, it will be noticed that
there is a distinct paucity of butterfly-eating birds in the wet
hill districts and that part of the coast subject to heavy rains ;
though mimicry occurs quite as commonly, if not more so, as
in the drier districts, where butterflies are less commonly met
with.

VALIDITY OF SOME FORMS OF MIMICRY. 735

Experiments on Wild Birds.

The following experiments were made on one bird in column 1,
two birds in column 2, and two in column 3 (see p. 724), a very
small percentage of the total number no doubt, but the best I
could do under the circumstances, and they indicate the lines
for future investigations.

Haperiments on Flycatchers at Nuwara Eliya, 1909.

The Indian Red-breasted Robin Flycatcher, Siphia hyperythra.

This bird is migratory and is found only in the hill districts ;
it is about the size of the European Spotted Flycatcher.

30.3.09. Deprived live Zerias hecabe one, Terias libythea one,
Hypolimnas bolina 3 one, Danais fumata one, Neptis leucothoé
one, and Appias galene one, of about two-thirds or three-quarters
of their wings, and put them on the ground near a tree from
which one of these birds was accustomed to feed. It first made off
with either the Veptis or Danais, I could not see which, then the
Appias. It then flew away, and J picked up one of the Terias
and Hypolimnas. This last had a good deal, perhaps half, the
wings left and fluttered about vigorously.

1.4.09. Lethe daretis one, Vanessa haronica one, Terias hecabe
one, and the same H. bolina, treated in the same manner as on
the last occasion and put in the same place. The bird carried off
the first two, but the H. Lolina seemed too large for it, as it was
for another Flycatcher which came along shortly after the first
had flown off. I could not see what became of the 7’. hecabe.
The same afternoon a similar experiment, but with no result.

9.4.09. This bird has evidently nugrated as I have seen none
since the last note. It nests in the Himalayas.

The Ceylonese Dusky-blue Flycatcher, Stoparola sordida.

This bird is peculiar to Ceylon but confined to the hill-tracts.

2.4.09. Placed Terias hecabe two dead, Argynnis hyperbius one,
Danais fumata three, Appias galene one, these latter alive but
largely deprived of their wings, near the cherry-tree much fre-
quented by Flyeatchers. The cock bird carried off one D. fumata,
its mate another ; the latter I was enabled to watch closely, and
it had extreme dithculty in swallowing the fly on account of the
wings. The former shortly afterwards returned and carried off
the remaining D. fumata, but the latter seemingly had had
enough of it. Shortly after a Red-breasted Flycatcher flew off
with the Appias. What became of A. hyperbius I do not know,
but the two dead Zerias were untouched.

Numerous specimens of the above butterflies were flying
about at the time, but I have never seen these Flycatchers molest
them.

5.4.09. Placed 7’. hecabe one, D. fumata one, and A. hyperbius

51*

736 LT.-COL, N. MANDERS ON THE

one, under the cherry-tree, having amputated both wings on
one side close to the body. The cock bird flew down almost
immediately and seized D. fuwmata, and as usual it had great
difficulty with the wings; about ten minutes afterwards it took
the A.hyperbius, but 7’. hecabe was left. I fancy the wings rather
‘‘ put off” the bird.

29.4.09. The birds have a nest with young in a cherry-tree
in the grounds. I put down 4. hyperbius 9, 7. hecabe, Neptis
lewcothoé, and one or two others; they had all been dead some
days and were very dry. After some time the bird noticed them,
flew down and seized the Argymnis, which had its wings closed
and showing the underside. It flew with it into a tree but very
shortly dropped it, it was evidently too dry. I found it had been
caught by the fore wings, one of which was gone with also a
portion of one hind wing. ‘The same afternoon put down live
Pyrameis cardui, N. lewcothoé, 7. hecabe, Appias nadina, and
Euplea core, but with both wings on oneside removed. The cock
bird flew down among them and caused a great flutter; 1t first
caught the Zerias, then the Neptis, and lastly the Huplea, which
provided a great chase. It carried them altogether to the nest,
but in feeding the young, the Huplwa escaped ; the bird was after
it in a flash, caught it again and carried it back to the nest. It
was very interesting to watch its efforts to get sucha large insect,
the size of our Camberwell Beauty, into the young one’s mouth.
Three or four times it had to take it out and manipulate it in its
own beak before another trial; eventually, it succeeded in forcing
it down the youngster’s gullet.

2.5.09. A half-winged live 4. hyperbius placed near the nest.
I am sure the bird noticed it, but beyond regarding it carefully
it did not molest it.

I am convinced from long and repeated observation that the
old birds never fed on butterflies themselves or fed their young
with them. <A critic, whose opinion I value highly, has objected
that because I never saw one of these birds capture a butterfly,
it is no proof that they did not do so and that very possibly
the difficulty of catching them would only induce pursuit
when the butterfly was off its guard and a capture possible.
I do not know why the birds should be more coy of capturing
a butterfly than a house-fly in my presence, and I can scarcely
believe they took the opportunity of my absence to do so.
Granted that difficulty of capture was the reason for non-
pursuit, what chance, it may be asked, would a young bird with
considerably feebler power of flight have of conducting a series
of tasting experiments on these butterflies? It is not infrequent
in the writings of advocates of mimicry to explain the rareness
of attack by difficulty of capture ; but by doing so they seemingly
forget that if such is the case with old birds, it makes tasting
experiments (with butterflies) very difficult for young ones.

VALIDITY OF SOME FORMS OF MIMICRY. 737

Haperiments on Birds in Colombo.
The Brown Shrike. anius cristatus.

28.12.08. Pinned a large Hypolimnas bolina 2 (a mimic of
HKupleeas) on a paling: in a few minutes the bird came along, and
directly it saw the butterfly it pounced upon it and carried it off,
and I could not see where it went, but I have no doubt it ate it.

29.12.08. Pinned a Hypolimnas misippus 2 on the paling.
The same Shrike saw it, seized it and held it in its claw, eating it
piecemeal and tearing off two or three wings. The following day
the same experiment was repeated with Danais chrysippus and
Telchinia viole, with the same result.

6.4.09. Nuwara Eliya. Noticed one of these birds, perched on a
twig, fly down and capture some insect on the ground. I watched
it for a long time, but though many butterflies flew past, it took no
notice of them. They comprised principally Argynnis hyperbius,
many Appias sp. ¢ Huplea core, Terias hecabe, and Lethe daretis.

28.11.09. Colombo. Put down Delias eucharis 2 , several Huplea
core, Danais limniace 2, Papilio hector, and Telchinia viole alive but
mutilated. A Shrike came and looked at them keenly from a tree
close by, but did not attack them. A small Cuckoo flew over
them twice, but took no notice of them.

21.10.09. Put down H. misippus $ 5, H. core 4. The Shrike,
perched in an oleander bush, evidently saw them, but for quite an
hour took no further notice though it took several insects close to
them. It eventually took one, perhaps two, Eupleas. I picked
up the others.

Magpie Robin. Copsychus saularis.

25.1.09. Put a number of Zerias hecabe (unpalatable) in the
porch of my house, where the Robin comes to feed morning and
evening; some had their wings removed, but it took no notice of
any of them.

4.2.09. Placed 1 7. viole, 1 FE. core, 2 T. hecabe, and 1 Precis
lemonias wingless, and 1 normal 7’. hecabe on the veranda. The
bird ate the Huplea with difficulty owing to its being very dry,
and it took the body of the wingless 7. hecabe in its bill, but
dropped it almost immediately as it was too dry ; it took no notice
of the others.

6.2.09. Placed specimens of the above on the veranda dead
but uninjured, and a wingless 7’. hecabe ; the bird ate the latter
but took no notice of the others.

Mr. Ormiston informs me that a Magpie Robin in his garden
has become so confidential as to take food from his fingers and
that it will eat ‘‘ almost any kind of butterfly when thrown to it,”
but he has never seen it catch one. Neither have I during the
seven years I have closely observed this species.

_ 21.10.09. Put down #. core 5 and D. chrysippus 1, with the
wings on one side removed. A young Magpie Robin, as shown by
its speckled breast, captured one Huplwa, and though evidently

738 LT.-COL. N. MANDERS ON THE

somewhat alarmed at its size, killed it and, after the usual difficulty
with the wings, swallowed it. It immediately captured and ate a
second, third, and fourth ; this last was a very vigorous insect and
fluttered a good deal before it succumbed. The bird was then
frightened and flew away, but carried the Huplea with it; it took
quite another ten minutes to get rid of the wings, and during
the process it twice flew off to capture small flies; it eventually
swallowed it. The D. chrysippus was, I have no doubt, eaten
by a Calotes which ran out of the grass close by where I had
put it.

22.10.09. Put down 2 Kupleeas, one dead and one moribund,
1 Hypolimnas bolina g dead with wings closed, and 3 H. mistp-
pus 3 alive and all lively. The young Robin immediately flew
down and tackled the Huplea, mangled it for some time and then
dropped it and flew away ; it returned shortly afterwards, picked
it up and flew away with it. Directly after, another young bird
flew off with and devoured the other Huplea. This attracted the
notice of the old birds, one of which, I think the cock, flew down,
but before he could seize a butterfly was hustled off by his mate,
who picked up two H. misippus and flew off with them. The
one H. misippus and H. bolina were left.

26.10.09. Put down H. misippus ¢ 2, H. bolina 2 1, Junonia
(Precis) almana 2, Pyrameis cardui and Catopsilia pyranthi, all
with wings on oneside removed. The H. misippus fluttered most
and attracted the attention of Calotes versicolor, which pounced on
and ate both of them; something then frightened it off. Next
an old cock Magpie Robin caught sight of the 1. bolina, seized and
killed it after a lively chase, and finally disposed of it. It was a
very long time beating off the wings and made many attempts to
swallow the fly, before it was finally successful. Three or four
times I thought the bird was going to leave it altogether. After
swallowing the fly, it went off and drank at the runnel close by.
Shortly afterwards, a young one of the same species caught sight
of the Junonia and captured it; this disturbed P. cardui close by,
and the bird dashed from one to the other, not able to make up
its mind which to take, when the old hen bird came and tried to
get one, but the youngster was too sharp for her and managed to
swallow both. I should have said that this bird hopped over the
Catopsilia, which was moribund and motionless, to seizethe Junonia.

Nore. I have now little doubt that so long as the butterfly is
motionless, resting, as these mutilated butterflies generally do,
with their wings expanded, they do not attract attention ; but
directly one moves, whatever the species happens to be, it is the one
to be seized and eaten, even though so-called palatable species are
close by. All these butterflies are flying commonly in the garden,
but I have never seen them molested.

28.10.09. Put down H. misippus 3 2, H. core 2, Catopsilia
pyranthi 2, Terias hecabe 1. The Robins came for them the
moment I went off; the old cock bird seized one Huplaa and made
off with it, and the young bird the other; this latter, after

VALIDITY OF SOME FORMS OF MIMICRY. 739

ineffectual efforts to break off the wings, left it and caught a
Catopsilia and ate it at once, and then the 1. misippus, afterwards
perching just above the place where I had put the butterflies.
The old bird saw the Huplea which had been killed by the young
one and regarded it for some time, then it flew down, pecked at
it, looked at it again and then flew off with it. The 7’. hecabe
managed to struggle into the grass and was lost.

Curiously enough, while this was going on, a Catopsilia
pyranthi was actually laying eggs within two feet of where I had
put down these butterflies, “and within twenty feet I found this
afternoon eges and larvee of Huplea on the oleander. This seems
to me to show that the butterflies, when whole, are not molested
because, I suppose, they are difficult to catch.

12.11.09. Put down three H. misippus 3, one a partial cripple,
the others with two wings off. An old hen Robin came at once,
and flew off with one of the wingless ones to a bush about twenty
yards off and ate it; but it did not seem very hungry. A young
one a few minutes after came and took the crippled fly and ate
it after the usual difficulty ; it came back in a minute or two and
ate the third one. The butterflies had emerged in the morning
and were consequently full of juice.

16.11.09. A young Robin made off at once with a newly emerged
H, nusippus 3.

18.11.09. Put down four mutilated recently emerged /7. misip-
pus $. The young Robin flew off with one from which the wings
had been removed and ate it; a few minutes after an adult cock
Robin came and ate the three others one after the other.

19.11.09. Put down four mutilated recently emerged H. misip-
pus 3S in the front garden on the drive. A Calotes ophiomachus
ran off with one anda Brown Shrike (Lanius cristatus) with two
others; the fourth, which had its wings only partially developed,
got into the grass, grew its wings, and eventually flew off. I
released three or four butterflies at the same time, and they flew
off strongly enough and were not chased by the Shrike, which was
sitting on a tree close by me.

21.11.09. Put down three H. misippus 2 form diocippus,
which resembles Danais chrysippus; the hen Robin came at once
and ate one and flew off with another ; a young bird followed its
mother, and flew off and ate the other. These butterflies had
hatched out that morning and the wings on one side had been cut
off. I next put down five more females, all with two exceptions
with the wings entirely removed. The cock bird took one of the
half-winged ones and then ate a wingless one. The young bird
then returned and finished off the remainder. ‘These female
butterflies evidently derived no protection from their resemblance
to D. chrysippus, and so far as two species of birds are concerned,
H. misippus is a palatable butterfly.

8.1.10. Found Lycana (Zesius) chrysomellus Q fluttering on
the ground ; it was headless and with a piece out of one hind
wing, probably caused by a Sparrow,

740 LT.-COL. N. MANDERS ON THE

During this month (November 1910) I have been breeding
Hypolimnas misippus freely, and the Magpie Robins come every
morning on the chance of getting one. I have tried them with both
males and females, crippled and perfect, and always with the same
result. They are immediately seized, well beaten, and swallowed
after considerable difficulty. 1 noticed on one occasion a perfect
female resting on the ground with wings widely expanded but
insufficiently strong to fly; the resemblance to D. chrysippus was
perfect, but the Robin seized it without hesitation.

One day the large Hill Crow—an occasional visitant—carried
off a crippled male with wings quite undeveloped ; it pecked at it
twice and then dropped it, shaking its head with every appearance
of disgust. I remembered that the insect had fallen into and was
well covered with the red liquid these butterflies always evacuate
on emergence, and thinking that this was the cause of the Crow’s
discomfiture, I covered a crippled female with the stuff and threw
it on the ground: a Magpie Robin soon came and saw it, and
shook its head once or twice after pecking at it, but it swallowed
it in a short time.

Mynah. Acridotheres tristis.

This bird belonged to Mr. O. Wickwar, F.E.S., who kindly
assisted me. The bird was quite young and was allowed perfect
liberty in a large garden, where it fed freely on grasshoppers
and other insects ; it had abundant insect food, and was also
accustomed to fill up its dietary by visits to the kitchen for
odd scraps. I have placed in brackets the presumed palatability
or otherwise of the species experimented on.

3.1.09. Given Huplea core (unpalatable), took it readily, but the
wings seemed to bother it considerably, so gave it another with
its wings shortened, this it ate readily enough and then went
back and finished off the first one. We then gave it Papilio lanke-
swara (palatable ?) which was also readily eaten. Half an hour
after gave Papilio (Menelaides) hector (unpalatable) ; this puzzled
it for a bit and it seemed disinclined to eat it, evidently on
account of the large wings, for when these were removed it ate
the body with relish, even hopping off the veranda after it when it
fell over the edge. <A couple of Telchinia viole (unpalatable) (an
Acreine) g and 2 followed, and then Hypolimnas bolina 2 (un-
palatable ?) and Delias euwcharis (unpalatable); all these received a
pinch on the head, were well pounded, wings partially removed,
and the remainder eaten.

24.1.09. The bird had been kept without food for some time
and was decidedly hungry. 3 P. hector, 1 7. viole, 1 Mycalesis
ceylonica (palatable ?), and 1 Polyommatus betica (palatable 2)
were put in a row outside the cage ; when this was opened the bird
hopped over them and made straight for the kitchen, where it was
accustomed to pick up odds and ends. After some persuasion it
ate 1 P. betica and 1 Lycena (Zesius) chrysomellus 2. Some
three hours afterwards it ate 1 Precis almana (unpalatable),

VALIDITY OF SOME FORMS OF MIMICRY. 741

1 Papilio hector and 1 P. aristolochic ; it seemed a good deal
worried by the wings. Afterwards neglected J/. ceylonica and
T. viole, but ate one wingless Huplea core.

The conclusion we arrived at from the above experiments was
that butterflies were not its natural food, but that when hungry
it would eat them indiscriminately, and that the palatability or
otherwise of butterflies was of no account with this species of

bird.

The dietary of Ceylon insectivorous birds is fairly well known,
and we are now in a position to discuss the questions—Do the
birds of this island eat butterflies largely ¢ If so, do they eat them
in sufficient quantity to produce any form of mimicry? and do
they show any discrimination in their attacks? In other words,
ean the terms palatable and unpalatable as applied to butterflies
be maintained.

As regards the first question, it will be granted that there is
a greater destruction of butterfly lfe than has hitherto been
supposed, and the following observation on a Bee-eater, though
necessarily a rough one, shows clearly that the destruction is
sufficiently severe to produce mimicry, provided of course that
the agents showed sufficient discrimination in their attacks.

The road from Trincomali on the north-east coast to Anuradha-
pura, runs through fifty-eight miles of thick forest which is cut
back some thirty paces on either side, thus affording a convenient
place for butterflies which avoid the dense jungle. Between the
hours of 9 A.M. and 10 a.m., I counted the number of butterflies
between the third and fourth milestones from Anuradhapura, and
they came to one hundred and ninety-five : the same day, in the
outskirts of the town I watched a Bee-eater feeding from 12.45
to 1.45 p.m., and during this time it caught twenty insects ; on
only one occasion could I be certain that the capture was a butter-
fly, and this was undoubtedly Catopsilia pyranthi. The bird feeds
from about eleven o’clock till five.

Motoring between these two places | calculated roughly that
there was a pair of these birds to the mile *, and consequently
the whole of the butterflies along this road would be cleared off
in about a fortnight unless they received an accession of strength.
The calculation is necessarily a rough one, but it gives a good
idea of the struggle for existence that is constantly going on.
This observation was made January 7th, 1909, at a time of year
when butterflies are less numerous than usual.

The question whether discrimination is shown by birds in their
attacks on butterflies is of the greatest importance in mimicry,
and on the answer depends the fate of both Batesian and
Miillerian mimicry.

I do not attach much importance to the fact, curious though it is,

* Bee-eaters are particularly fond of perching on telegraph-wires.

742 LT.-COL. N. MANDERS ON THE

that in the observations I have been able to collect, the Hupleines
and Danaines, popularly supposed to be highly distasteful, figure
more largely as victims than any other group. I believe this to
be simply due to the fact that these butterflies occur in very
large numbers, and not that distinct preference is shown for
them. Admitting that more evidence is needed, I doubt whether
future investigations will reveal any marked preference in those
birds which are mainly instrumental in the destruction of
butterflies, for the reason that their dietary is of such a mixed
character ; and if this were so, or if what I have here set forth
be considered sufficient to settle the question, it is difficult to
avoid the conclusion that the unpalatability of these butterflies
has been assumed on insufficient data. It is interesting to recall
Professor Meldola’s remarks written so long ago as 1879, when
Miller first propounded his theory of mimicry (Proc. Zool. Soc.
Lond. 1879) :—

“ ...it may be fairly asked how far we know that such
imitated groups as Heliconius, Huplwa, Danais, Acrea, etc., are
distasteful. But very few observations have, as far as | am
aware, been made even upon these groups which are generally
admitted to be the objects of imitation, and I certainly know of
no systematic exper iments conducted with these models and their
insectivorous foes.”

The Bee-eaters seem to show some partiality for the yellow and
white butterflies of the Catopsilia and Appias group ; but whether
this is more apparent than real is not clear. It may be that these
butterflies are more readily seen and easier to capture than
others ; but if it could be proved that there isa distinct preference
for them, it is noteworthy, considering the destruction that
undoubtedly takes place, that though very variable they do not
act as models or mimics, or form Miillerian combinations, either
in India or Ceylon.

Failing the butterfly-eaters, what evidence is there that the
birds of group 2, and group 3, show preference in their more or
less desultory attacks ? There is no doubt that those experimented
on showed none, and that they took no notice of butterflies unless
they were mutilated and rendered easy of capture. I should
much wish to see further experiments on wild birds of these two
groups undertaken, but if the butterfly-eaters do not conduce to
mimicry, it is doubtful to my mind if the partial feeders would
do so.

In the present state of our knowledge it is difticult to say what,
is or what is not an unpalatable genus, and the position is further
complicated by the proposition that unpalatable species are killed
in numbers sufficient to produce a special form of mimicry. It
is unfortunate that theoretical considerations rather than observa-
tions and experiments in the field have hitherto preponderated
in this matter. It seems to me that the terms palatable and
unpalatable are not justified at present,

VALIDITY OF SOME FORMS OF MIMICRY. 743

The Miillerian Theory.

The supporters of the Millerian theory hold the view that it is
chiefly by the attacks of the young inexperienced birds that this
form of mimicry is produced. Professor Poulton puts the case
as follows :—‘ The Miillerian theory presupposes that only young
birds test the palatability of a few members of each convergent
group in their locality and henceforward, except when driven by
hunger, avoid all the members, so that the recent tendency to
explain so many of the resemblances on Miillerian rather than
on Batesian lines is in harmony with the conclusion that the
members of such groups are not greatly attacked by adult birds.”
(Hssays on Evolution, p. 270.)

I have already expressed the opinion that it is unlikely that
young birds, except those in group 1, indulge in tasting experi-
ments on butterflies, but as IT am quite willing to admit that such
an opinion may be founded on insufficient data, and as I was un-
able to find the necessary evidence required by the Miillerian
theory, I approached the subject by another line of investigation,
which depends on the time of the nesting of the birds and the
broods of the butterflies.

The birds breed once a year, not twice as is the casein Mauritius.
They begin in March or April, sometimes early in May, according
to the season. When the March or April rains known as the
little monsoon bring out a large increase of insect life, the birds
immediately begin nesting amt the young birds are off the nest
and begin to forage for themselves i in May, June or early July.
The average life of an insectivorous bird is probably not more
than four or five years, and we may assume that tasting experi-
ments gradually grow fewer in number and are completed when
the bird is about six months old, 2. e. about the month of October.

In estimating the number of broods of butterflies in the year,
which vary much according to the species, I will direct attention
to two of the more striking cases of mimicry, that of the Kuplcas,
forming a Miillerian combination, and Papilio polytes with its
trimorphic female mimicking P. aristolochic and P. (Menelaides)
hector. They may be taken together. In January, February and
March, that is to say in the dry weather (I am speaking more
particularly of the plains), there is a very small but continuous
series of broods which depend on the weather for their develop-
ment. If it is very dry, the eggs, larvee or pups, as the case may
be, lie dormant, but with favourable meteorological conditions
such as a shower of rain, the eggs hatch, the larve shake off their
lethargy and feed, or the butterfly emerges. Mr. Mackwood in-
formed me that on March 24, 1908, in “his garden at Colombo,
eggs, larve and pupe of Euplea core could be found together on
the same tree. The majority of the pupe do not, however, hatch
out but remain quiescent until the April rains, when there is an
astonishing outburst of butterfly and other insect hfe. With the
onset of the south-west monsoon ut the end of May or beginning

744 LY.-COL. N. MANDERS ON THE

of June, the broods become larger in numbers and more frequent,
and this goes on until the end of August or beginning of
September, when there is a further spell of dry weather similar to
but not so pronounced as that in the early part of the year, when
the broods again become smaller and less frequent, but at the same
time produce the individuals which take part in the migratory
flights of the monsoon in November and December.

Strictly speaking, P. hector and P. aristolochic, though following
the above sequence of events, do not usually form part of the
flights, but they are nevertheless at their maximum at this time ;
the Eupleas and Polytes undoubtedly do so. We have now to
judge what influence the inexperienced young birds off the nest
in May, and their experiments concluded in October, can have on
these species. A butterfly the size of Huplwa core pairs during
its first flight, if we may judge by the cabinet condition of those
ovipositing, sail begins to lay its eggs three daysafterwards. The
usual number is about two hundred and fifty, which are deposited
according to the weather in about ten days (I have known one
hundred egos laid in five days). What becomes of the parent
after this? Whether she dies a natural death or becomes the
victim of a tasting experiment is immaterial, her time of danger
is a brief fortnight. As the females are less in evidence than the
males, fewer of them would be captured, especially if we agree with
Professor Poulton’s opinion that the Miillerian theory presupposes
that only young birds test the palatability of a few meinbers of
each convergent group in thei locality. To bring to such per-
fection the cases of mimicry I have selected, we must assume that
such a victim would be one having less converging characteristics
than the others ; and it must also be borne in mind that unless
she is killed within three days of her emergence, she will have laid
a certain number of eggs which will produce butterflies similar to
herself. It is difficult to understand how the broods of butterflies,
numbering some thousands of individuals, born between October
and the following nesting season, would be in any way affected
except in the very smallest manner. Nodoubt Nature is infinitely
slow in her methods, and we have no reason to suppose that these
cases of mimicry have been produced, otherwise than by a very
lengthy process of weeding out ; but even if we grant this, there
is a still greater difficulty in the case of Hypolimnas misippus, the
well known mimic of Danais chrysippus. In Ceylon the former
appears on the wing in October, when as I have said tasting
experiments are over. It remains on the wing until the end of
the year, when it disappears until the following autumn. There
are so far as I can see only two ways of getting over this diffi-
culty—either by assuming that the inherited tendency to produce
this form of mimicry has become so fixed that the withdrawal of .
the factor that produced it is immaterial, though there is no reason
for this supposition, or that there is a more or less constant influx
of the species from India. There is very little doubt that a certain
number of Ceylon butterflies in their annual migratory flights

VALIDITY OF SOME FORMS OF MIMICRY. 145

find their way to India, but I can find no evidence of a reverse
condition of things, and one can scarcely credit that the few chance
stragglers which possibly find their way across the sea could keep
up this perfection of mimicry in the south of the island. The
supposition that the sudden appearance of a previously unknown
species would produce further tasting experiments will not hold
good in this case, as the mimic so “closely resembles its model
D. chrysippus, which is on the wing all the year round.

Huperiments on Young Lirds wr Confinement.

I am extremely doubtful as to any real value accruing from ex-
periments on caged birds, whether nestlings or adult. No one, I
imagine, believes that all butterflies taste alike: no doubt some
are more tasty than others, and caged birds fed upon butterflies
even with other insect food would no doubt learn in time to dis-
tinguish the different kinds ; but this procedure to my mind begs
the question, as 1t assumes that butterflies are an ordinary article
of food in the wild state, a proposition regarding which the
evidence here brought forward does notaltogether support. The
ease 1s different with Coleoptera, Hemiptera, Diptera and the
like, which are known to be the staple food of birds. Lloyd
Morgan’s carefully conducted experiments leave no doubt that
certain species of birds, probably all, have very little instinct as to
what is good, and what not, and that they learn by imitation
and tasting experiments. My observations lead me to believe
that the former is very important. I briefly epitomize my own
conclusions.

1, Young birds probably learn at first in a general way
what is their natural provender by what is brought to them
in the nest.

2. That this is further developed when they have left the nest
but are too weak to accompany the parents when they are
foraging for food.

3. That when they accompany the parents, as they do for a
longer or shorter time according to the species, they notice the
insects caught and attempt to capture them themselves.

4. When they are left to shift for themselves they carry on
what they have learnt, and during this time they undertake
tasting experiments, but with the exception of the birds in group 1,
those on butterflies are few in number ; first, because they have
rarely or never had butterflies brought to them in the nest ;
secondly, because they have very rarely seen their parents catching
them, and so neglect them; thirdly, because they have considerable
difficulty i in catching them, and the process of getting rid of the
wings is tedious and lengthy and the morsel fluffy, and possibly
not always agreeable. If these observations are confirmed by
further experience, they would account for the fact that attacks
on butterflies are less frequent than those on other insects.

746 LT.-COL. N. MANDERS ON THE

Instances of Imitation by Young Crows.

July, 1910. 1 saw an old crow and two young ones on the Rifle
Green this morning; one of the youngsters had hold of a bone
with a piece of gristle attached to it. It was so firmly adherent
that the bird could not detach it as the bone constantly moved
with the bird’s efforts, and eventually it gave up. Then the old
bird, which had been standing by all the time, went to the bone,
put its foot on it, thereby gaining a purchase, and tore off the
gristle without difficulty; the young bird after two or three
attempts did the same.

September, 1910. An old crow had a piece of hard boiled
potato off which it was picking pieces and giving them to a
full-fledged young one close by. A goodly number of detached
pieces lay on the ground and attracted the attention of some
other crows, which flew down and began picking them up ; seeing
this the youngster did likewise, though it made no attempt to do
so before their arrival.

Seasonal Dimorphism—Cryptic Defence.

I should not conclude this study of mimicry without discussing
that form of it which is known as ‘‘ cryptic defence,” and especi-
ally that which is so noticeable in the seasonal changes of so
many tropical butterflies.

It is commonly believed to have been produced by natural
selection acting through the medium of insectivorous foes, the
more exact and perfect imitations found in the dry season being
due to the paucity of insect life at that time of the year, which
produces a greater keenness in pursuit and a greater struggle for
existence. The argument has been put forward in full by
Professor Poulton in his ‘ Essays on Evolution,’ page 203.

I hope in the near future to deal more exhaustively with this
subject, but at present will only direct attention to two species
occurring in these islands, a study of which does not favour the
usually accepted views. The contention for the production of the
dry season form rests upon the premiss that ‘the dry season is a
time of far greater pressure than the wet”; for although the
enemies of insects are fewer, the insects themselves are pro-
portionately even more reduced, and ‘‘the light thrown by recent
investigation leads us confidently to believe that the differences
between the seasonal forms—hitherto devoid of interpretation—
have a meaning and a value in the struggle for existence and
came into being under the sway of natural selection ” (Powlton).

Though it is probably correct to say that in countries such as
S_ Africa and India, which havea continental climate, the seasons
are such as to produce a wealth or poverty of insect life, it is de-
cidedly incorrect to assume the same with regard to the islands
we have been investigating. There is no doubt that at no season
of the year is there in any of them a paucity of insect life, and at

VALIDITY OF SOME FORMS OF MIMICRY. 47

no time would an insectivorous bird, or reptile, find any difficulty
in procuring its daily sustenance.

Bourbon and Mauritius are very largely under sugar cultivation,
and this necessitates constant manuring of the “fields! with a
consequent abundance or super abundance of flies of all sorts
throughout the year. The rainfall, though greater in the wet
season, 1s not infrequent during the dry season, and this also
favours insect life; and if we add to this the consideration of
the practical absence of butterfly-eating birds and reptiles, we
can estimate the difticulty of believing that these changes are the
outcome of natural selection in these two islands.

In Ceylon Mr. E. Ernest Green, who has lived thirty years in
the island and who knows every part of it, writes as follows :—

‘Though insects are more abundant at certain seasons, J have
never experienced any part of Ceylon where there was anything
approaching a dearth of them. I know that Iam always busy
pinning and setting throughout the year.

‘““T sometimes wish that pee was a short dead season, when one
could devote oneself to other work without being distracted by the
constant accumulation of material.

“T doubt if Melanitis is ever subject to much worry from birds.
It lives in the shade and never moves during the daytime, unless
flushed by some big animal. Jam now receiving (16.8.10) both
dry and wet season forms of J. tambra from Kandy.”

An allied species Melanitis leda occurs also in Bourbon and
Mauritius, and it is to be remarked that the dry season forms
begin to appear before the advent of the dry season, that is to say
before any form of stress would tend to make itself felt.

In explanation, it may be suggested that the butterfly was in-
troduced from the locality where natural selection produced these
changes and that it is simply carrying on an inherited tendency.
That it is an introduced species is highly probable, but it has been
known to entomologists in Bourbon and Mauritius for at least
sixty years, and it differs in no way now than in the time of
Boisduval. It is difficult to believe that the factor which pro-
duced this cryptic defence being removed and no longer required
would not have led to some other form of colouring, or a return
to that ancestral type from which these forms were evolved. The
above remarks apply equally to Wycalesis narcissus, Precis rhadama
(introduced 1858), and Terias floricola, and I have made a further
study of Terias hecabe in Ceylon. It is very frequently the case
that the wet form continues to appear well into the dry weather
and vice versa, but to a less extent; this has been remarked on
frequently, but so far as I know no exact observations have been
made. In Colombo there was no rain from November 19th till
December 10th, 1908, thence to January 6th, 1909,-70 of an inch,
but of this no less than ‘57 fell on one day (Dec. 19th); such an
absence of rain in a tropical country at once causes a general drying
up of vegetation and the assumption of dry weather conditions.
At weekly intervals I captured all the Terias I could, which were

748 ON THE VALIDITY OF SOME FORMS OF MIMICRY.

accustomed to breed on a hedge of Madras thorn in an isolated
position in my garden; the results of such counts showed 73 wet
forms, 1 intermediate, and 19 dry, and it was not until January
28th that the wet forms were entirely replaced.

The butterfly takes exactly a fortnight to pass from larva to
imago, and thus we have approximately three or four broods of
wet season forms produced under diy season conditions. We
must assume that as the butterfly is seasonally dimorphic it stands
in need of protection, yet so far as I could ascertain the wet forms
suffered no diminution though exposed to what were, or should
have been, adverse circumstances.

The butterflies rested during the night and in cloudy weather
on the under surfaces of the leaves of Vincéasp.?, a small flowering
shrub with pink flowers and small oval green or frequently yellow
faded leaves. It often collected gregariously, two or even three
being on the same leaf and perhaps ten on the same plant. The
position was an admirable one for protection from the wet, and
also from small predatory foes which seldom look upwards when
hunting for prey. The appearance of the plant is the same
throughout the year, and the butterfly derived no advantage from
its change from one seasonal form to the other.

The following experiment makes me still further doubtful of
these effects being due to natural selection.

If we take the pupa of a somewhat similarly coloured butterfly
which is not seasonally dimorphic, such as Papilio demodocus or
Papilio demoleus, and expose it to a hot dry temperature, we can
produce an insect with much of the yellow on the under surface
replaced by red. I am almost persuaded that these rusty red
spots are a vestige of a character at one time common to certain
Pierines and Papilionines which is more or less reproduced by
heat and dryness, if of sufficient duration and intensity under
natural conditions in the Pierines, but in the Papilionines oniy
under artificial stimulation of a like but exaggerated character.

CoNCLUSIONS.

1. It has been shown that in Bourbon and Mauritius there are
no butterfly-eating birds or reptiles; so that the cases of mimicry
occurring there cannot be due to their influence.

2. In Ceylon it has been conclusively shown that the butterfly-
eating reptiles are impartial feeders.

3. That a trained observer can distinguish the majority of
these mimetic butterflies at a distance of about twenty or thirty
feet and frequently at the same number of yards; and this being
so, it is certain that a bird which has to depend for its existence
on its powers of observation, could after a few failures be able to
discriminate them at the same and probably at a considerable
greater distance.

4. That Drongos feed largely upon Eupleeas, and this being so,

DISTRIBUTION OF THE GENUS MEGAPODIUS IN PACIFIC. 749

a Papilio mimicking them obtains no protection in the vicinity of
these birds.

5. There is no bird in Ceylon known to eat butterflies that dis-
tinctly discriminates as an adult between one species of butterfly
and another.

6. It has been shown that there is a great destruction of
butterfly life in the dry zone, and that here, if anywhere, Miillerian
or Batesian mimicry might be induced, but the destroyers are
largely migratory and their attacks are not selective.

7. That the number of broods of butterflies which occur be-
tween the termination of tasting experiments in one year and the
commencement of them in the next is so great that any influence
which could be wrought by such is almost inappreciable.

8. The little evidence available shows that young Ceylon birds
imitate their parents in their choice of food; but as regards butter-
flies, the fact that there is no discrimination shown by adults leads
one to conclude either that few or no tasting experiments were
undertaken in youth, or, what is more probable, that their taste
with regard to them is indifferent.

9. Itis questionable, and so far as an accurate knowledge of one
species goes it is definitely shown, that that form of mimicry re-
presented by wet and dry season forms (cryptic defence) is not
produced for the protection of the species, inasmuch as many
(four) succeeding broods of the wet weather form may be found
under dry season conditions without detriment to the species.

34. The Distribution of the Avian Genus Megapodius in the
Pacific Islands. By J. J. Lister, M.A., F.R.S., F.L.S.,

¥.ZS.
[Received and Read May 9, 1911. |

(Text-figure 166.)

The Megapodiide or Mound Builders are, as is well known,
large birds, with comparatively feeble powers of flight, con-
stituting a family of the order Galline. They are distributed over
the islands of the Kast Indian Archipelago and Western Pacific,
from the Philippines and Borneo to the New Hebrides, and are
found in several parts of the continent of Australia. Four out-
lying species of the genus J/egapodius are found in the Nicobar,
the Pelew, and the Marianne Islands, and, far out in the Pacific,
on the little island of Niuafou, belonging to the Tongan group.

As we cannot suppose that the birds found in these outlying
islands, remote from the other species, can have flown across the
intervening tracts of ocean, we are presented with the problem:
How did they reach these islands ?

The solution to which M. Oustalet gives his adhesion, in his

Proc. Zoot. Soc.—1911, No. LIT, 52

750 MR. J. J. LISTER ON THE DISTRIBUTION OF THE

monograph on the Family *, is that all these localities have been
at one time connected by land, which has since been to a large
extent submerged.

*Kermadec I$

“o
a

| rat Howell
Map illustrating the Distribution of the Genus Megapodius.

NEW ZEAL
160

Text-fig. 166.

ica

ep}
ayes
a
a <
= =|
2
She
Ry

Wallace had, in 1876 7, expressed the opinion that the Megapode

* “Monographie des Oiseaux de la famille des Mégapodiidés.” Ann. d. Sciences
naturelles, 6 sér. t. 10 & 11, 1880-81.
+ Geographical Distribution of Animals. London, vol. 11. p. 342.

AVIAN GENUS MEGAPODIUS IN THE PACIFIC. 751

of the Nicobar Islands had probably been introduced by the Malays,
but Oustalet cannot regard this explanation as plausible, on the
ground that ‘“‘ we possess no positive proof of the domestication of
Megapodes by the Malays or the savage people inhabiting the
Oceanic Islands.”

My object in this paper is to point out the reasons which appear
to me to make it probable that the distribution of the genus
Megapodius has been, as Wallace suggested in the case of the
Nicobar bird, considerably modified by human agency ; and that
the species found in these outlying Pacific islands have been
carried there by man.

Since Oustalet published his monograph the Megapodiide have
been again reviewed by Ogilvie-Grant in his Catalogue of the Game
Birds in the British Museum*; and in 1901 Rothschild and
Hartert t gave their revision of a portion of the genus Megapodius.

The determination of the limits of the species of this
genus is difficult on account of the variation in size and colour
presented by the birds inhabiting the same locality, and the fact
that the characters of those from different localities often merge
into one another. On comparing the results arrived at by the
authors mentioned, we find that nineteen species were enumerated
by Oustalet. If we take from these W. wallacei, which has been
placed by Ogilvie-Grant in a separate genus (Hulipoa), and M.
brenchleyt and M. brazieri, which had then been described only
from eggs or young birds, there remain sixteen species. Five of
these are united in two species by Ogilvie-Grant, and seven which
inhabit the area dealt with by Rothschild and Hartert are allowed
by these authors only subspecific rank under two specific names.
On the other hand, the species J. macgillivrayi, which is united
by Oustalet with the widely extended WZ. duperreyi, and regarded
as only a subspecies by Rothschild and Hartert, is reckoned a
distinct species by Ogilvie-Grant.

The peculiar nesting habits of the Megapodiide are well known.
Most of the species scrape together large mounds of earth or sand,
with or without vegetable matter, and the female deposits her
egos, which are very large for the size of the bird, at intervals of
several days, in excavations in these mounds. Incubation is
effected by the heat of the slowly fermenting mass, aided by that
of the sun, or by the sun alone; and the young are hatched in an
advanced state of plumage. They receive no attention from their
parents, and in some cases at least they are able to fly on the day
on which they are hatched. In some species many pairs of birds
frequent the same mounds or laying-grounds.

In almost all the countries where Megapodes occur, their large
eggs are highly valued by the natives as food, and their laying-
places are frequently visited for the purpose of obtaining them.

* Catalogue, Vol. xxii. 1893. Also‘ Game Birds’: Allen’s Naturalists’ Library,

London, 1895-7. :
+ “Notes on Papuan Birds.” Novitates Zoologicx, vol. viii. p. 135, 1901.
| 5o*

52 MR. J. J. LISTER ON THE DISTRIBUTION OF THE
There is clear evidence that the birds are at least semi-
domesticated in some localities.

Mr. C. M. Woodford *, speaking of J. brenchleyi (AL. eremita
of B. M. Catalogue) on Guadaleanar, in the Solomon Jslands, says :
“The birds lay in open sandy clearings, generally near the sea,
which are kept clear of shrubs and undergrowth by the natives,
and by the sand being constantly turned over by the birds... .
Many ‘thousands of birds congreg: ate at the same place, the laying-
yards being often some acres in extent.” Of the little island of
Savo, to the north of Guadaleanar, the same author writes } :
“although only about the size of a large pigeon ’ * the megapode
‘lays an ege bigger than that of a duck,” and “eggs form an
important item in the daily food-supply of the natives.” ‘The
megapodes lay their eggs on two large cleared sandy spaces and
nowhere else on the island. Upon these no weeds or grass can
grow as the ground is constantly being turned over by the birds
when digging ¢ holes to lay their eggs, and by the natives when in
search of them. The sandy spaces are fenced off in plots which
belong to different owners.” He adds that the natives are quite
indifferent as to the condition of the eggs when they eat them, it
is all the same to them whether they are newly laid or well
advanced towards hatching.

Mr. John Brazier, writing of a collection of eggs of Megapodes
exhibited before the Society: says t, ‘‘ When at San Christoval ”
(in the Solomon Islands) “I was shown an egg that Perry, a
white man living there these last five years, said was laid by the
‘Wild Fowl, and upon my visiting him a few days later, he had
just obtained another from the nest: of his domestic fowls.”

M. Freycinet, in his narrative of the voyage of the Uranie §, says
of the species which was discovered by this expedition on the
Marianne Islands: ‘‘ Espéce de gallinacée de couloir noir que jadis
les anciens Mariannais élevoient auprés de leurs cabanes ; elle est
aujourd’hui fort rare. Nos naturalistes lui ont donné le nom de
Mégapode la Perouse.”

In the volume on Zoology (p. 125) of the same work the
naturalists Quoy and Gaimard say of JIL. freycineti, which they
discovered on the island of Waigiou, to the west of New Guinea,
and which is now known to occur from the Moluccas to Western
New Guinea: ‘“ Sur les iles Vaigiou et Boni, ces oiseaux parois-
sent vivre dans une demi-domesticité, d-peu-prés comme les
canards qui habitent les marais que traverse la petite riviére de
Sévre, (Charente Inferieure).” One, brought by the natives, lived
sever: al days on the Uranie.

Professor J. Stanley Gardiner, whose investigations of the
Fauna and Flora of the Maldive and Laceadive Islands are well

* The Naturalist among the Head-hunters, PP. 100-101.
P. Z.S. 1888, pp. 249 & 260.
P.Z.S. 1874, p. 607.

Bi age autour du Monde..... sur les corvettes de S.M. PUranie et la

AVIAN GENUS MEGAPODIUS IN THE PACIFIC. 753

known, informs me that a sultan of the Maldive Islands, who
died in 1878, introduced Megapodes into an islet covered with
cocoanuts and scrub, forming part of the great atoll of Male in
that Archipelago. Where these were obtained is not known, but
Professor Gardiner thinks it probable that they were imported
from the Nicobar Islands, between which group and the Maldives
there is regular trade communication.

We have thus from four widely separated localities definite
evidence of the more or less complete domestication of Megapodes
by the natives. Finally Guillemard *, referring to Wallace’s view
that the Nicobar Island bird was introduced, says ‘‘ that this is
not impossible must be evident to every traveller in the Malay
Archipelago, for birds of this genus are often seen in captivity.”

With regard to the powers of flight of Megapodes, they are
compared by some authors with those of barn-door poultry.
Oustalet, however, recalls the fact t that a young specimen of
M. freycineti flew on board ‘ La Coquille,’ with a favouring breeze,
when that vessel was “plus de deux milles” (over two miles)
from land.

Le Souef ¢ mentions that IJ. duperreyi, although the birds are
‘very poor fliers,” occurs on the scrub-covered islands ‘a good
many miles” from the N.E. coast of Queensland. He surmises
that they may have been blown out during cyclones. .

Finsch § speaks of IW. senex, the species inhabiting the Pelew
Islands, as occurring on nearly all the sandy and rocky islands of
the group. Some of these are separated by intervals of some
three or four miles. He considers that the bird, ‘‘ which is a good
flier” (the term is of course used in a relative sense), may oc-
casionally fly from one island to the other. He also mentions
that the eggs are systematically taken by the natives.

But when all allowance is made for their powers of flight, it
would seem an extravagant suggestion, and one which I think
has never been made, that Megapodes could by this means have
reached the outlying islands in which they are now found. The
Pelew Islands are separated by nearly five hundred miles from the
Philippines, the nearest land to the west, and by a rather greater
distance from New Guinea to the south. The Marianne Islands
are some 600 miles to the E.N.E. of the Pelew Islands. Niuafou
is nearly 1000 miles to the east of the New Hebrides, the nearest
islands to it on which a species of Megapode exists.

We may now examine the geological nature and some other
conditions of these outlying islands in the Pacific on which Mega-
podes are found, as well as the characters of the species living
on them.

The island of Niuafou, or New Hope Island, although politi-
cally part of the Tonga group, is situated almost halfway between

* Cruise of the Marchesa, vol. ii. p. 122 (footnote), 1886.
+: IG. c, vols Xi. p69 ay
+ Ibis, 1899, p. 16.
§ “Die Vogel der Palau-Gruppe.” Journal des Museum Godeffroy, Heft viii.
Bd. iii.) 1875, p. 30 (p. 162 of the volume).

754 MR. J. J. LISTER ON THE DISTRIBUTION OF THE

the Fiji aud Samoa Islands. It is described in Findlay’s South
Pacific Directory * as a voleanic island with black lava rocks all
round the shores. It is 3 to 34 miles across, well wooded, and
some 500 to 600 ft. high. In the centre isa brackish-water lake
at sea-level in which are hot springs. Friedlander +, who visited
the. island in 1897, describes his visit to the nesting places of the
birds, which are on the shores of this lake. He had to swim
round some of the rocky points to reach them, and found the
temperature of the water that of a warm bath, and the rocks
under water too hot, in places, to rest his hands on them. He
says the island is an intermittently active crater, largely composed
of basaltic rock. An eruption occurred in 1886, when the whole
island was covered with ashes, and the Megapodes were nearly
exterminated. Owing to the ‘tabu’ imposed by the chief their
numbers had increased again, so that there was a fair number at
the time of his visit. The birds do not build mounds, as do many
of their congeners, but lay in holes which they excavate in the
voleanic sand. He is inclined to attribute the heat of the sand,
which he found on digging for the eggs, to the volcanic action.
We may note in passing that P. & F. Sarasin } found that the
Maleo (Aegacephalon maleo) of Celebes lays in the neighbourhood
of hot springs, as well as on the sea-shore.

The birds of Niuafou were collected by Mr. F. Hiibner and
described by Dr. Finsch §. With the exception of the Megapode
all belong to common Tongan species, but Finsch remarks on the
absence from the collection of four species (Ptilotis carunculata,
Halcyon sacra, Lalage maculosa, and Colluricinela heinei) which
besides being common in Tonga are present, or represented by
allied species, in Samoa and Fiji. At first sight 1t would appear
that the absence of these species from Niuafou might be at-
tributed to the destructive eruptions of the volcano, but as these
species are unrecorded from the not very distant group of Uvea
(Wallis Id.), their absence from Niuafou cannot certainly be
attributed to that cause. It must be admitted, however, that a
small volcano, still in intermittent activity, is the last place on
which the remnants of an ancient fauna would be expected to
survive. Had the bird been found on the high and ancient land
masses of Fiji or Samoa, the case against this view would not
have been so strong, but the birds of these islands have now been
so fully collected as to make it in the highest degree unlikely
that so large and useful a bird as a Megapode should have been
overlooked in them.

The Niuafou species, Megapodius pritchardi, was described by
the late Sir Walter Buller in his Supplement || to the ‘ Birds of
New Zealand,’ and included, under the name of ‘The Southern

* 3rd edition, p. 558.
+ Ueber die Nestlicher d. Megapodius pritchardii auf der Insel Niuafou.”
Ornithologische Monatsberichte, vil. p. 87, Berlin, 1899.
t Zeits. d. Gesellschaft f. Erdkunde, Berlin, 1894, pp. 375, 388, 396 & 398.
Z.S . 1877, p. 782.
|| Vol. i. p. 31.

AVIAN GENUS MEGAPODIUS IN THE PACIFIC. 755

Megapode,” in that fauna—on what appear to be wholly in-
adequate grounds.

In 1887, Mr. T. F. Cheeseman, the well-known Curator of
the Auckland Museum, to whose knowledge and kindness many
visitors to New Zealand ave indebted, visited the Kermadec
Islands, which are a scattered group lying nearly halfway between
the North Island of New Zealand and the Tonga Islands to the
N.N.E., and some 400 or 500 miles from either. Mr. Cheeseman
reported * that a Mr. Johnson, who had resided on Sunday
Island (a volcanic island, the most northerly of the group) about
fifteen years before, told him that “ prior to the eruption of 1876
a bird inhabited the floor of the large crater, which made
mounds of sand and decayed leaves, two to three feet high,
laying its eggs in the mounds. He was in the habit of visiting
the mounds for the sake of the eggs and young birds, and has
frequently taken 5 or 6 of the latter from the same nest at one
time.” The eruption of 1876 covered the floor of the crater and
apparently killed out the species. Mr. Cheeseman cautiously
observes that the evidence, such as it is, seems to point to the
former existence of a species of Megapodius on this island.

We may remark that the statement that five or six young
birds were taken from the same nest at one time is hardly
in accordance with the habits of the genus, for the eggs being
laid at some intervals the young ones are not of the same age,
and leave the mounds to feed for themselves soon after they are
hatched. The statement would be more appropriate to the young
of the Grey Duck (Anas superciliosa) which frequents this island.
It is the mound-building habit which, as Mr. Cheeseman says,
“seems to point” to the existence of a Megapode on Sunday
Island.

Sir Walter Buller + on the strength of this evidence includes
Megapodius pritchardi among the birds of New Zealand, in which
region the Kermadec Islands are included. He says: “1 have
no doubt whatever in my mind—notwithstanding the apparent
difference in their nesting habits—that Mr. Cheeseman was right
in his conjecture” that the Sunday Id. and Niuafou birds were
identical. (It will be noted that Mr. Cheeseman conjectured that
the genus, not the species, was identical.) On the discrepancy
that whereas the Niuafou bird lays in burrows, the Sunday Id.
bird is stated to have built mounds of sand and leaves two to three
feet high, he remarks (p. 33): “If the latter observation was
accurate it may have been due to circumstances of locality ana
environment, and by no means negatives the assumption of these
birds being one and the same species.” As we have seen, the
statement that the Sunday Island bird built mounds is the only
evidence we have of the existence of a Megapode on that island.

Through the kindness of Mr. Ogilvie-Grant I have had the good

* “Qn the Birds of the Kermadec Islands.” Trans. and Proc. of the N. Zealand
Institute, vol. xxiii. (1890) p. 219.
+ Supplement to the Birds of New Zealand, vol. i. (1905) p. 31.

156 MR. J. J. LISTER ON THE DISTRIBUTION OF THE

fortune to meet at the Natural History Museum Mr. Iredale, who
has recently resided on the Kermadecs with the object of studying
their fauna. He assures me that he was not able to obtain
any confirmation of the report of the existence of a Megapode
on Sunday Island which was given to Mr. Cheeseman, and finds
that the successors of his informant are not inclined to regard
that report as worthy of very serious consideration.

It therefore seems to me that we have no good evidence that
the genus Megapodius formerly inhabited the Kermadec Islands
and absolutely none that JZ. pritchardi lived there.

One good result, however, we owe to Sir Walter Buller’s
enthusiasm in claiming this species as a member of the New
Zealand fauna, and that is a plate representing the bird in a
condition of plumage not hitherto figured.

M. pritchardi belongs to the section of the genus with the
back and upper surface of the wings rufous brown, the breast
and belly lead or slaty grey, and in its general coloration perhaps
is nearest J/. cumingi, Dillw., of the Philippines and Borneo.
In the type specimen described and figured by G. R. Gray *,
and now in a somewhat dilapidated condition in the British
Museum, the bases of the quill-feathers, except the first, are
white; there is also some white among the upper tail-coverts.
A specimen in the Leyden Museum was described by Schlegel +
(and I have had an opportunity of examining it) which has,
as he observes, the upper tail-coverts pure white. The first
specimen which came to the Auckland Museum was described by
Buller ¢. It had no white on either quill-feathers or tail-coverts,
though the rectrices were white at the base§. As it was not
known then that this skin came from the same locality as
M. pritchardi, and it differed so considerably from the type of that
species, it was described by Buller as a new species—W. huttonz.

Buller’s plate (pl. ii.) in the Supplement to the ‘ Birds of
New Zealand’ shows no white in the plumage. In the descrip-
tion he says (p. 32):—“ Although absent in this specimen ”
(that named M. hattoni and perhaps the specimen figured in the
plate) ‘‘ most examples have a patch of white covering the basal
parts of the primaries and secondaries, the extent varying in
almost every individual. Some also have white markings on the
upper tail-coverts and basal part of the tail-feathers.”

It appears then that the feature by which M. pritchardi stands
apart from all other species of the genus—the occurrence of white
at the bases of the primaries and elsewhere—is a varying and
inconstant character.

To return to the comparison with MZ. cumingi, I find that
M. pritchardi has the top of the head slaty-brown rather than
brown, the sides of the head rather paler, the mantle brown tinged
with slate rather than olive-brown, and the belly a paler brown.

* P.Z.S. 1864, p. 41, pl. vi. + Mus. Pays-Bas, vii. p. 64.
Transactions of the N. Zealand Institute, vol. iii. (1870) p. 14.
Hutton, Trans. N. Zealand Institute, iv. (1871) p. 165.

AVIAN GENUS MEGAPODIUS IN THE PACIFIC. rey I

We may note that the species inhabiting the New Hebrides,
4M. layardi, belongs to the group of species having the mantle
and upper parts blackish grey, uot rufous brown as in
M. pritchardi.

There is one other fact which seems to point to the view
that the Niuafou bird is not indigenous to that island, viz., that
the native name, Mallow, is the same as that applied to several
species of Megapodiide i in the Malay Archipelago. This appears
in the specific name of Megacephalon maleo of Celebes. Mega-
podius cumingi is called moleo kitjil (= little moleo) by the
natives of that island, in distinction, probably, from the larger
Megacephalon maleo*. M. layardi is the Malow of the natives s of
the New Hebrides. Onustalet says that this name appears to be
applied indifferently to Megapodes by Malay hunters. Tis occur-
rence on Niuafon, far out in the Pacific among a population of
Polynesian speech, seems to suggest strongly that at some time
the name arrived at the island with the bird.

With regard to the Marianne and Pelew Islands, I have less
evidence to bring forward, but it was in the former group that
Quoy and Gaimard were told that the Megapode (i. laperousti)
had been domesticated. It is so closely similar to Uf. senex from
the Pelew Islands that M. Oustalet has regarded them as of the
same species T.

The geological structure of the Pelew Islands is discussed by
Semper 7, who shows that the islands are composed in part of
raised coral, in part of voleanic rock, formed during submarine
eruptions. If this is the case, there can be no remains on the
Pelew Islands of the fauna of a subsided land-mass, supposing
such a mass to have existed.

From the description of the Marianne Islands in the account
of the Voyage of the Uranie, above quoted, it is stated (Historique,
T. 2, p. 253) that they seem to have been formed in the remote
past by submarine eruptions, which have raised the floor of the
ocean, and that the reefs which have formed about the islands as
they have risen above the waves have since been raised with them.
So that it would appear that the same remark is applicable to
these as to the Pelew Islands.

4M. laperoustiand M.senex have the upper parts blackish grey, as
have M. freycineti, from the Moluccas and New Guinea, Jf. geel-
vinkianus from New Guinea and some adjacent islands, and
M.layard: from the New Hebrides. But they differ from these
and other species in the french-grey colour of the feathers of the
head. We must conclude therefore that this character has been
developed since their isolation, or else that the parent stock has
either not yet been discovered, or has become extinct.

In the British Museum Catalogue three “doubtful species”

* Meyer & Wiglesworth, Birds of Celebes, vol. ii. 1898, p. 671.
+ Of clic Guan Alea aes Library : Game Birds, vol. u. p- 182.
= The Natural Conditions Enxistenee as they affect Animal Life, by Karl

of
Semper, Chapter 8: International Scientific Series, vol. xxxi. London 1883, pp. 234
264.

758 DISTRIBUTION OF THE GENUS MEGAPODIUS IN PACIFIC.

are mentioned (p. 446) by Ogilvie-Grant, viz.: 1. Young birds
said to have been obtained on Lord Howe’s Island; 2. The
Megapode of Sunday Island in the Kermadec group; and
3. M.¢ andersoni of Gray from New Caledonia. The first of these
Mr. Ogilvie-Grant assures me is now known to have come not
from Lord Howe’s Island, but from New Hope Island, another
name for Niuafou. With the second I have already dealt. The
third is based on a reference in the MS. of Anderson, who
accompanied Cook’s third voyage, to a bird he called Tetrao
australis and briefly described as follows :—“ fusca nigraque ;
pedibus nudis.” The subsequent exploration of New Caledonia
has not revealed the presence of a Megapode on that island.

To sum up: There is evidence of the domestication or semi-
domestication of Megapodes in several parts of the area they
inhabit,—viz., in the Solomon Islands, Western New Guinea and
the Marianne Islands, and of their introduction into the Maldive
Islands. There is no satisfactory evidence that a Megapode has
ever existed on any Pacific island east of a line bordering the
Philippines, Solomon Islands, and New Hebrides except the
Pelews, Marianne Islands, and Niuafou. The geological character
of these islands, so far as we know it, lends no support to the
view that they could preserve the fauna of a sunken land-mass.
The birds of the Pelew and Marianne Islands are almost identical,
and on the latter group they were domesticated. The bird of
Niuafou is called by a Malay name.

When we consider the complex movements of the races of the
Western Pacific, of which there is much anthropological evidence,
and how easily Megapodes might be introduced into a new locality
by a canoe provisioned with their eggs, which are a staple native
food, it would appear that we have in human agency a probable
key to some of the anomalies of their distribution.

The analogy of the distribution by native agency of domestic
fowls, cousins of the Megapodiide, and of the dogs and pigs
which were found by the early voyagers on the Pacific Islands, is
obvious. :

How far the same cause may have been operative within
the main area occupied by the genus, and have given rise to the
anomalies in the distribution of the species alluded to by Oustalet,
is too large and complex a subject for me to attempt to deal with.

The Savo natives, says Mr. Woodford,* speaking of Jf. eremita
of the Solomon Islands, have a curious legend connected with this
bird. They hold the Shark in great veneration and say that their
island was made by the Shark, who brought stones together and
placed upon them a man, a woman, the Yam plant, and the
Megapodes. Things went well for a time and the people increased
and so did the Megapodes. At last the people went to the Shark
and complained that the Megapodes made havoc among the yam

* C. M. Woodford, P. Z. S. 1888, p. 249.

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ON THE MORPHOLOGY OF THE HELICINID#. 759

patches by digging holes to lay their eggs; so they asked the
Shark to take the Megapodes away. This was done, but now the
natives missed the Megapodes’ eggs, so they asked the Shark to
bring the Megapodes back but to confine them to one spot.
This request was also complied with, and the result may now be
seen. The Megapodes lay their eggs in two large and broad
sandy spaces, and nowhere else on the ccbnd.

I suspect that there is more than a grain of true history in
this legend, and that it records the fact that when the ancestors
of the natives came to the island, they brought with them two
main staples of their food-supply—yams and Megapodes.

35. Contributions to the Morphology of the Group Neritacea
of the Aspidobranch Gastropods.—Part Il. The Hett-
cinip#. By Givpert C. Bourne, M.A., D.Sc., F.R.S

F.Z.S.
[Received April 29, 1911: Read May 9, 1911.]

(Plates XX X.-XLIL*)

When, two years ago, the Society published the first part of my
contributions to the mor phology of the Neritacea (2), I had already
accumulated a number of observations on the anatomy of the
Helicinide, but deferred the publication of them until I was able
to obtain specimens of different species from various parts of
the Pacific region. Having experienced considerable difficulty in
obtaining specimens sufficiently well preserved for microscopical
examination, the publication of my results has been long delayed,
with the result that I lose the claim to priority for several minor
discoveries concerning the anatomical features of this family, for,
in the meantime, Thiele (10) has given an account of the anatomy
of Hydrocena cattaroensis in which is included a description of
the female generative organs of Helicina hubaryi, and the following
deser iptions lose much of the novelty they would have possessed
had they been published as soon as the facts were ascertained.

Previous to the publication of Thiele’s paper, our knowledge of
the anatomy of the Helicinide rested, for the most part, on
Isenkrahe’s (4) account of the anatomy of Helicina titanica.
Isenkrahe gave a sufficiently accurate description of the external
anatomy, the muscular system, the greater part of the alimentary
tract, and the pulmonary cavity, but he failed altogether to
distinguish the kidney, and his descriptions of the heart, the
nervous system, and the reproductive organs are defective. These
imperfections notwithstanding, Isenkrahe was able to confirm
Troschel’s opinion that Helicina, on account of its rhipidoglossate
dentition and other anatomical characters, was closely related to
the Neritide. .

Von Jhering (5) in 1877 placed the Helicinacea and Proserpinacea

* For explanation of the Plates see pp. 806-809.

760 PROF. G. C. BOURNE ON THE

in his class Orthoneura, order Rostrifera, sub-order Rhipidoglossa,
and gave a fairly accurate description, unaccompanied by a figure,
of the nervous system of Helicina (Stwranya Wagner) beryllina
Gld. E. L. Bouvier (3), in his great work on the nervous system
of Prosobranch Gastropods, gave a very complete account of the
nervous system of H. sagraiana d’Orb. and H. brasiliensis Gray,
laying stress on its close resemblance to the nervous system of the
Neritidz, and in addition he made some further observations on
the general anatomy, partly confirming and partly correcting and
adding to Isenkrahe’s descriptions. In 1902 Thiele (9) described
the male generative organs of Helicina japonica, and last year
he gave a description with a diagram of the female organs of
H. kubaryi, in addition to a succinct but sutticiently exhaustive
account of the general anatomy of Hydrocena cattaroensis.

The geographical distribution of the Helicinide, as is well
known, presents several interesting and ditticult problems. By
far the greater number of species are insular and confined to the
tropics. Such species as are found on continents are for the
most part limited to regions near the coast, very few being known
to occur any considerable distance inland. No Helicinide are
recorded from Africa. In Europe the group is represented only
by the genus Hydrocena from the Dalmatian coast, and this genus,
as Thiele’s recent work has shown, differs in several important
anatomical characters from Helicina and its more closely allied
genera. Georissa,a subgenus of Proserpina, is the only represen-
tative of the group in India, and no Helicinide have as yet been
recorded from Ceylon. The number of genera and species reaches
its maximum in the Antilles. The genus Helicina, as restricted
by Wagner, is fairly abundant in Mexico and the Centrai
American republics, and extends northwards into Texas and
Florida, southwards into Ecuador and Peru on the west coast and
to the south of Brazil on the east coast of S. America. Few
species, however, are recorded from the Pacific coast of S. America,
but, notwithstanding their comparative rarity on these shores,
the group reappears in great abundance in the Pacific islands,
extending as far east as the Marquesas and Paumotu Islands, and
having many representatives in the Society, Samoan, Friendly
and Fiji Islands, and in the New Hebrides and New Caledonia.
Several species occur on the east coast of Australia, and some
few are recorded from New Guinea, Celebes, Borneo, and Sumatra ;
none, so far as I can ascertain, from Java. But in this part of
the world the Helicinide attain their maximum in the Philippine
Islands, which are only second to the Antilles in the number of
species. From thence the group extends north, beyond the
tropical zone, to the Bonin Islandsand Japan. A few species
are found beyond the south-east coast of China and Siam, others
again in the Malay Peninsulaand Burma. Several species are
found in the Andaman and Nicobar Islands, but the group is
very poorly represented in the Indian Ocean. Aphanoconia
(Helicina) theobaldiana G. & H. Nevill is recorded from the

A

MORPHOLOGY OF THE HELICINID&. 761

Seychelles, and Psewdotrochatella undulata Morelet is a subfossil
form from Mauritius. None is known from Madagascar.

In a recent work of great value to the student of geographical
distribution A. J. Wagner (12) has revised the family Helicinide,
and, founding his diagnoses chiefly on the characters of the
operculum, has broken up the old genus Helicina into no less
than thirteen genera, reserving Lamarck’s appellation for the
American and Antillean forms which conform to the original
definition of the genus. Of the remainder I mention the largest
genera. Sulfurina has its centre in the Philippines and extends
thence to the Andamans, Nicobars, Moluccas, New Guinea, and
Tahiti. Aphanoconia, which also seems to be centred in the
Philippines, extends widely, to Japan, S$. China, the Malay
Archipelago, the Andamans, Nicobars, Seychelles, Moluccas, and
through Micronesia and Melanesia to the Paumotu and Sandwich
Islands. Stwranya has its centre in Fiji and Tonga, and extends
thence to the Carolines, Sandwich, Society, Hervey, and Solomon
Islands. Orobophana is found in Queensland and N.S. Wales and
extends through nearly the whole of Polynesia. Palcohelicina.,
with its subgenus Ceratopoma, is again a Philippine genus, and
extends to New Guinea, the Bismarck Archipelago, the Solomon,
Louisiade and Pelew Islands. The last-named genus is, according
to Wagner, closely allied to Helicina sensu restricto. Again, the
subgenus Letorquata of Helicina, which occurs in Mexico and
Central America with outliers in Florida and Texas, affords,
according to the same author, a transition to such a characteristic
Antillean genus as Alcadia. A consideration of these statements
leads to the conclusion that the Helicinidee are capable, by what
means we know not, of wide dispersal across seas and oceans, and
find conditions most suitable to their existence in proximity to
the sea. They appear to have originated in Mexico and Central
America, and to have spread eastwards to the Antilles, where
they found the conditions specially suitable, and have been
differentiated into several genera (Aleadia, L ucidella, Eutroch atella,
Priotrochatella, Proserpina) and numerous species, and one species
(H, substriata convexa Pf.) has found its way to the Bermudas.
Others have extended down the eastern coast of S. America, but
the Atlantic Ocean has proved an impassable barrier to their
further extension eastward. On the Pacific side the group has
been transported by some means unknown to us to the Pacific
Islands, and it would appear from the evidence that it did not at
first effect a lodgment in the more eastern islands, but in the
Philippines, from which centre it has spread in all directions—
eastward throughout Polynesia and to the Sandwich Islands,
southward to New Guinea and Australia, northward to Japan

and China, westward through the Dutch ‘Indies and Malaya to
the Andaman and Nicobar’ Islands. Very few have traversed
the Indian Ocean to reach the Seychelles and Mauritius.

Very little is known of the geological history of the group.
Helicina occurs in the post-Pliocene of N. America, but the

762 PROF. G. C. BOURNE ON THE

ancestral forms must have lived at a much earlier period, for
Proserpina is recorded from the Kocene of the Isle of Wight, and,
according to Kobelt (6), shells referable to the same genus have
been found along with Helia, Planorbis, Vi alvata, and three
species of Veritina inthe Lias of Somerset. There is some reason
for suspecting the correctness of the identifications in the last
case, and [ am unable to find any corroborative evidence of the
occurrence of Proserpina in the Eocene, but the distribution of
the Hydrocenids points to a geological history reaching well
‘ back into Tertiary times. Dawsoniella from the Carboniferous of
Illinois has been attributed to the Helicinide, but I have already,
in the first section of this memoir, discussed the affinities of this
genus and pointed out that it must be a case of convergence.

However this may be, paleontology throws very little light on
the origin and distribution of existing Helicinide, and rhe I
began this work I hoped, not only to give a full description of the
anatomy of a typical member of the family, but also, by the
comparison of the anatomy of Pacific and West Indian forms, to
discover some clue to the distribution of the group with its two
main centres in the Antillesand the Philippines. In this, as will
appear, I have been disappointed. From whatever part of the
world they may come, the anatomy of the different species and
even genera of Helicinidz is so closely similar that it is hard to
find any difference between them. It is true that I have not been
able to procure many species of Pacific Helicinide, but I have
examined fairly well preserved specimens of Or -obophana, Aphano-
conia, and Paleohelicina, and these three genera may be taken as
typical of the more widely distributed Pacific forms.

The material at my disposal was as follows :—

I. Antillean forms.

Alcadia palliata Ads. Contrivence, Walderston, Jamaica.
Alcadia hollandi Ads. Swing Hill, Walderston, Jamaica.
Lucidella aureola Fér. Bog Walk, Spanish Town, Jamaica.
Eutrochatella pulchella Gray. Bog Walk, Spanish Town,

Jamaica.

The above were kindly collected for me and preserved in
Perenyi’s fuid by Mrs. G. B. Longstaff, F.L.S.

Il. Pacific, Australian, and Indian forms.

Aphanoconia gouldiana Forbes, from Torres Straits: for
specimens of this species I am again indebted to Mrs.
Longstaff, who procured them for me from Mr. C. Hedley,
of the N.S. Wales Museum.

Aphanoconia andamanica Benson.
Aphanoconia merguiensis Pfeiffer.
Aphanoconia rogersii, sp. n.

These three species are from the British Museum and formed
part of the collection made in the Andaman Islands by Mr. G. Rogers.

MORPHOLOGY OF THE HELICINID®. 763

They were numbered respectively 16, 30,and 31. The first two I
have identified without difficulty, the third appears to be new
to science, and I will give a diagnosis of it in the latter part of
this paper. Iam indebted to Mr. KE. A. Smith for these and for
the two following species :—

Orobophana pachystoma ponsonbyi Smith. Admiralty Islands.
Paleohelicina ide Wagn. Amboina.

In describing the anatomy it will be convenient to take Alcadia
as the type, and to note such differences as may exist between it
and the other genera at the end of the description of each system
of organs.

External Characters, Mantle, Mantle-cavity,
and Muscular System.

Isenkrahe (4) has given an account of these so sufficient and
accurate that it is not necessary for me to do more than call
attention to some special features exhibited in fig. 1 (Pl. XXX.),
which is a representation of a left side view of Alcadia palliata :
the mantle has been cut through on the left side close above the
columellar muscle, the cut has been extended back to nearly the
extreme hinder end of the mantle-cavity, and the mantle has been
turned over towards the right. As compared with the Neritide,
in this and in all the other species of Helicinide that I have
examined the foot is attached to the head and body by a longer
and narrower. pedicle, the opercular lobe is relatively smaller, the
snout is narrower and longer, the columellar muscles of greater
antero-posterior length, and the whole body is longer, giving the
appearance of an increase in the coiling of the visceral mass, but
this last feature is more apparent than real, as I shall show.
A glance at the figure shows that the increased length of the
body is chiefly due to the elongation of the post-tentacular region
and the part of the body immediately following. Using
Amaudrut’s (1) phrases, we have an almost extreme case of
“allongement posttentaculaire,” followed by an “allongement
dorsal,” and many of the peculiar features of helicinid anatomy
are'to be explained by the excessive growth in length of these two
regions. The post-tentacular region lies above the anterior two-
thirds of the columellar muscle, and its posterior limit is marked
by two or three deep wrinkles of the body-wall. The body-wall
of this post-tentacular region is fairly stout and muscular, and
the epidermis is, as a rule, deeply pigmented. The colour differs
in different species. It is nearly black in Alcadia, grey shading
posteriorly into white in Hutrochatella pulchella, yellowish grey
in Lucidella aureola, a dark chocolate-brown in Paleohelicina ide,
and a bright chestnut-brown in Aphanoconia gouldiana. In the
post-tentacular region are contained the buccal bulb and the
greater part of the cesophageal pouches. In the dorsal region
following on the post-tentacular the body-wall is thin and nearly

764 PROF. G. C. BOURNE ON THE

transparent, the musculature is feeble, and the epidermis is not
pigmented, This dorsal region is relatively of considerable length ;
its concave lower border corresponds very closely in length with
the surface of insertion of the left columellar muscle; its upper
surface extends back to the pericardium. It contains nearly all
the coils of the intestine, the cesophagus, the radular sac, and the
hinder lobes of the ceesophageal pouches. Its roof forms the floor
of the hinder part of the mantle-cavity. In consequence of the
elongation of these two regions, but particularly of the dorsal
region, the mantle-cavity is continued very far back ; so far that,
measured from its most anterior to its most posterior limit, it
makes nearly a complete turn of a spiral, whereas in Verita and
Neritina it makes little more than half a turn. Broad in front
where its roof passes from the right to the left columellar muscle,
the mantle-cavity becomes narrower and narrower posteriorly and
ends ina pointed cul-de-sac below and somewhat to the right of the
lower surface of the visceral mass. Its extreme posterior limit is
not quite visible in fig. 1. With the hinder end of the mantle-
cavity the pericardium has also been carried very far back. It is
laid open in fig. 1 to show the position of the heart. It will be
seen that the single auricle seems to lie behind the ventricle, and
not in front of it as does the larger left auricle in the Neritide. In
horizontal sections, such as those depicted on Pl. XX XIII. figs. 18
and 19, this posterior position of the auricle forces itself so much
upon one’s attention that I was led to form the theory that the
single auricle of the Helicinide corresponds not to the left and
larger, but to the right and rudimentary auricle of the Neritidee ;
and in my memoir on the morphology of the latter family (2,
p. 833) I prematurely gave expression to this view, which seemed
to me the more probable because I found that in correlation with
an increase of the pallial vessels in Septaria the right auricle
became relatively larger and took an obviously larger share in
carrying blood from the pallial vessels to the ventricle. But since
then, after a careful study of the relative positions of the kidney,
the uropore, the rectum, and the heart in the Helicinide, I have
satisfied myself that this view was erroneous.

As a consequence of the dorsal elongation of the body all
these organs have been rotated through an angle of rather more
than 90°, in such wise that the pyloric end of the stomach, which
in the Neritide is directed forward and to the left, comes to lie
at the right posterior end of the body in the Helicinide, and
the swollen esophageal end of the stomach, which is posterior
and somewhat to the left in the Neritide, is directed anteriorly
and to the right in the Helicinide, forming a conspicuous
rounded prominence at the extreme end of the visceral mass
(Pl. XXX. fig. 1, St.'). To understand the nature of this rotatory
movement the reader should refer to fig. 42 of my memoir on
the Neritide. This figure represents a horizontal section of
Paranerita gagates, and shows the position of the cesophageal (St.)
and pyloric (St.') divisions of the stomach. It should be noticed

MORPHOLOGY OF THE HELICINIDA. 765

that the extremity of the visceral mass, lying to the right, is
wholly occupied by the liver and gonads. If, now, the reader will
lay a tobacco-pipe in front of him on the table, the bowl (repre-
senting the cesophageal division of the stomach) to his right, the
stem (representing the pyloric division of the stomach) to his left
front, it will occupy much the same position as does the stomach
in the figure referred to. If, while the bowl is kept pressed
against the same spot on the table, the stem is lifted up and
rotated through an angle of rather more than 90° till it points
over the observer's right shoulder, the whole pipe will have been
rotated through an angle which brings it into the position of the
stomach of the Helicinidea—the animal being supposed to be
placed foot downwards upon the table with its head turned away
from the observer. As all the organs of the left side of the
body, including the posterior end of the mantle-cavity, the peri-
cardium, the heart, the kidney, and the coils of the intestine,
have shared in this movement of the pyloric end of the stomach,
their positions have been nearly completely reversed, and the
left auricle instead of lying in front of the ventricle has come
to lie behind. The right auricle has entirely disappeared in the
Helicinide, and the rectum, having undergone some degree of
displacement in connection with the above-described movement
of rotation, is no longer enveloped by the ventricle.

The kidney has also undergone a curious and at first sight a
puzzling change of position. In the Neritide, as described in
my previous memoir (2, see Pl. XLVI. fig. 1 for its position in
Septaria, Pl. LIV. fig. 29 for its position in Paranerita gagates),
its glandular part lies to the left hand of and partly below the
rectum : posteriorly the glandular part opens into the spacious
non-glandular bladder or ureter, and the latter runs forward,
below the glandular part, to open by the uropore into the mantle-
cavity on the right of and close to the base of the ctenidium.
In this family the greater part of the kidney lies in the roof
of the mantle; it is only its posterior extremity that passes below
the rectum and invades the visceral mass, where it lies just above
the pyloric end of the stomach. The effect of the rotation of the
last-named organ in the Helicinide is that the kidney has been
carried round till it comes to le wholly in the visceral mass, on the
lower side of the latter, between the loop of the rectum which
passes round this region and the pyloric division of the stomach,
as may be seen in the series of sections (Pl. XX XIII. figs. 17 to 20)
and in the diagram (Pl. XXXIV. fig. 24), which is a reconstruction
from this series of sections. The kidney, in short, has been
turned completely round, so that its originally posterior end looks
to the left front and the uropore opens into the right hinder
corner of the mantle-cayity, the reno-pericardial canal, main-
taining its relation to the uropore, into the right posterior corner
of the pericardium. It is further to be observed that the visceral
mass, though apparently more coiled, is really less coiled in the
Helicinidee than in the Neritide. In all systematic werks stress

Proc, Zoou. Soc.—1911, No. LITI. 53

766 PROF. G. C. BOURNE ON THE

is laid on the fact that the internal partitions of the shell are
absorbed in both these families. This absorption has not pro-
ceeded quite so far in the Neritide as in the Helicinide. In
Nerita and Paranerita there is a recess in the upper right-hand
part of the shell which contains that lobe of the visceral mass which
consists wholly of liver and gonads and represents the visceral
spire of other Gastropods. This recess and the lobe of the
visceral sac corresponding to it are not found in the Helicinida :
the wall of the cesophageal division of the stomach comes very
near to the surface (Pl. XX XIII. figs. 17 & 18), and the liver and
gonads are disposed at the sides of and above the pyloric division
of the stomach. The more coiled appearance of the whole is
due to the elongation of the post-tentacular and dorsal regions,
not to the retention of a larger section of the visceral spire of the
presumed gastropod ancestor than in the Neritide.

From what precedes, it follows that most of the peculiarities of
the Helicinid organization are the result of excessive growth and
elongation of a particular region of the body, and it is an
interesting confirmation of the correctness of the above account
of the manner in which the Helicinid organization has been
derived from the Neritid that, if one makes a plasticine model
of the stomach, kidney, rectum, and intestinal coils as they occur
in Paranerita, and then rotates the stomach in the manner
described, the intestinal coils assume very nearly the position
found, with more or less variation in detail, in all Helicinids.

After this general explanation of the mutual relations of the
principal visceral organs in the Helicinide, I need only refer to
particular features in the several systems of organs which I have
to describe in detail. Before proceeding, it should be put on
record that there is not a rudiment of the ctenidium in the
Helicinide, and I cannot even find a trace of an osphradium.
The cephalic penis, characteristic of the males of the Neritide, is
also absent, and there is no externa] difference between the males
and females in any of the species that I have examined.

®

The Alimentary Tract.

The complex of organs formed by the buccal cavity, the pharynx,
the cesophagus with its smaller and larger glandular annexes, the
radula, the radular sac, and the odontophoral cartilages and their
muscles, can only be studied by dissection, and this is by no means
an easy task in animals so small as most species of Helicinide are.
The relations of the various organs to one another are far too
complicated to trace out in sections. The following description
applies chiefly to Alcadia palliata and A. holland, but will serve
almost equally well for any of the other species that I examined,
for all are very much alike except for the details of the radular
teeth.

The mouth is a gaping circular orifice, situated at the extremity
of the downturned snout: it is surrounded by folded muscular lips.

MORPHOLOGY OF THE HELICINIDA. 167

The buccal cavity occupies the snout, in front of the tentacles.
It is a simple funnel-shaped cavity bounded by a rather thick
muscular wall, the internal surface of which is thrown into about
19 or 20 longitudinal folds. The cavity is lined by a layer of
rather long columnar epithelial cells which secrete a thick cuticle.
Dorsally the buccal cavity is prolonged backward into a little
glandular diverticulum which lies above the median part of the
cerebral commissure. ‘The buecal cavity is separated from the
pharynx by a constriction, deepest on the dorsal side, where the
cerebral commissure lies in it. In a surface view, before dis-
turbance of the various parts, this constriction is not visible
from above, as it is covered over by the anterior salivary glands
shortly to be described, and muscle-fibres pass from the walls of
these glands to the walls of the buccal cavity and of the snout.
Consequently the cerebral commissure seems to be embedded in
the buccal mass.

The passage from the buccal cavity to the pharynx is narrow.
The pharynx is a relatively spacious sac, of which the cavity
is continued posteriorly into the esophagus above and into the
radular sac below. Beneath and at the sides of the anterior
end of the radular sac lie the odontophoral cartilages, the
anterior ends of which project forward into and occupy the
greater part of the lower moiety of the pharyngeal cavity. It
will readily be understood that, in consequence of the projec-
tion of the anterior ends of the odontophoral cartilages into
the pharyngeal cavity, the latter extends round them both at
the sides and below. Below the cartilages the pharyngeal
extension forms a broad flattened diverticulum, reaching back
nearly to the posterior ends of the anterior cartilages, as far
as the point marked a in fig. 3 (Pl. XXX.). Laterally, the line
of attachment of the pharyngeal wall to the anterior odonto-
phoral cartilages is roughly indicated in the same figure by the
curved line running upward and forward from the point « towards
the opening of the esophagus. It results from this arrangement
that in an oblique section, such as is represented in fig. 4
(Pl. XXX.), the pharynx appears to give off two posterior diver-
ticula, lying outside the anterior ends of the odontophoral
cartilages. The inner walls of these diverticula are thin and
composed of a single layer of cubical epithelial cells: they are
continued round the anterior and upper edges of the cartilages
into the lining membrane of the radular sac. The outer wall of
each diverticulum is strengthened by a thin plate of cartilage,
too small and transparent to be recognized in dissection, but
readily recognizable in sections. These lateral pharyngeal carti-
lages serve for the attachment of muscles, one set of which run
forward to be inserted on the walls of the snout, the other set
run backward and are inserted on the odontophoral cartilages ;
the former are protractors, the latter retractors, of the walls of the
pharynx.

A portion of the epithelial lining of the outer wall of each

53*

768 PROF. G. C. BOURNE ON THE

diverticulum is composed of very long attenuated epithelial cells,
among which are long club-shaped glandular cells. This glandular
strip may be traced upward and forward to the thickened lip of
the esophageal opening, where it forms a prominent ridge
which passes above into the anterior pair of salivary glands, to
be described shortly.

As is shown in P]. XXX. fig. 2 and in the section drawn in
fig. 4, the radular sac opens into the pharynx by a widely gaping
aperture situated in the trough-shaped depression between the
anterior ends of the anterior odontophoral cartilages. Posteriorly
the radular sac passes between the posterior odontophoral cartilages
and runs at first to the right of and below the cesophagus, but
soon mounts upwards and to the left, the two organs, oesophagus
and radular sac, being twisted round one another as shown in
fig. 2. The radular sac is short in Alcadia palliata and is of no
great length in any of the species that Ihave studied. The greater
or less length of the radular sac appears to be an individual rather
than a specific character. The characters of the radular teeth will
be dealt with in a separate section.

The odontophoral cartilages were described in some detail by
Isenkrahe for Helicina titanica, and I have but little to add to his
account, A ventral view of these structures in Alcadia palliata
is given in Pl. XXXI. fig. 5, and a sketch of a dorsal view of the
same structures in Hutrochatella pulchella in fig. 6: both figures are
drawn to the same scale. As may be seen from the specimens
figured, the odontophoral cartilages exhibit specific differences in
relative size and proportion, but these differences are of too slight
and elusive a character to be expressed in a description, and scarcely
important enough to make it worth while to give aseparate figure
for each:species examined. The essential structure is the same
in all. There are two pairs of cartilages, an anterior and a
posterior. Each member of the anterior pair is a plate having
the form of a more or less elongated isosceles triangle; the
margins of the plate ave thickened and rounded, the central
portion remains thin. The plate is bent in such a way that its
lower margin is bent inward posteriorly and its upper margin
outward. The posterior margin or base of the triangle, forming
the articular surface for the posterior cartilage, runs obliquely
from above downwards and inwards. ‘The lower margins of the
two cartilages are connected by a tough fibrous band, and their
thickened edges serve for the attachment of the intrinsic and
extrinsic odontophoral muscles. The posterior cartilages are
short conical masses; their apices directed backwards; their
ventral surfaces convex and their dorsal surfaces more or less
concave. Their broad anterior endsare shaped to correspond with
the articular surfaces of the anterior cartilages, and the two are
firmly held together by muscular fibres, whose arrangement is
indicated in fig. 5. It follows from the above description that
the odontophoral cartilages form the sloping sides of a V-shaped
trough, the concavity of which looks upwards and supports the

MORPHOLOGY OF THE HELICINID®. 769

anterior part of the broad radular ribbon. The median and
admedian radular teeth lie in the floor of this trough, the great
lateral teeth he in the angles between its floor and sides, and
the uncini form curved rows running upwards and backwards
along its sloping sides. The radular ribbon is attached by strong
muscular bands to the cartilages. These muscles run obliquely
forwards from the radula to be attached to the anterior cartilage
of either side, and obliquely backwards to be attached to the
posterior cartilages, these two sets of muscles causing the ribbon
to slide forward and backward over the smooth surfaces of the
cartilages.

The relation of the cesophagus to the radular sac and odonto-
phore is shown in fig. 3, which is a drawing of a dissection of
these structures in Alcadia palliata. The left cesophageal pouch
and the left side of the cesophagus have been cut away, and the
roof of the cesophagus has been lifted back to the right side to
show the entrance to the right cesophageal pouch and other
structures. In the angle of the deep fold between the cesophagus
and radular sac are seen the buccal ganglia (g.bwe.) lying just above
the middle of the anterior odontophoral cartilages. ‘The opening
of the cesophagus into the pharynx, situated just in front of and
above the buccal ganglia, is irregularly funnel-shaped, with thick-
ened and folded lips projecting forward into the pharyngeal
cavity. laterally, these lips are deeply grooved, and on either
side the groove is continued backward and downward into the
lateral pharyngeal diverticulum described above, and upward and
somewhat forward into the anterior salivary gland of its own
side. These anterior salivary glands are formed by a pair of pocket-
like forward projections of the cesophagus, which in their natural
position lie side by side and form a pair of pouches lying above
the cerebral commissure. Between them is an anterior cecal
diverticulum continuous behind with the median dorsal groove of
the anterior part of the csophagus. When separated by an
incision in the mid-dorsal line and turned outwards, the anterior
salivary glands present the appearance shown in fig. 2. Inter-
nally their walls are raised into a number of thick glandular
ridges: the outermost of these ridges is specially thick and is
continued downward, in the groove passing to the side of the
cesophageal orifice, into the glandular ridge on the outer wall of
the pharyngeal diverticulum, as has been described above.

The anterior section of the cesophagus, lying above the
odontophore, is fairly wide. Internally, its walls present a
number of longitudinal glandular ridges, and in the mid-dorsal
line there is a deep groove bounded internally by prominent
ridges; posteriorly this groove shallows and eventually dies out.
On either side of this anterior section of the esophagus is a
gaping oval orifice (figs. 2 & 3, 0.@.p.) leading into the large
cesophageal pouches, or, as some would call them, the posterior
salivary glands. The last-named structures are capacious irregu-
larly lobulated sacs with large cavities. Their inner walls are

770 PROF. G. C. BOURNE ON THE

lined throughout by a glandular epithelium, consisting, as far as
T was able to Gbeeenne! almost wholly of very long goblet-shaped
secretory cells containing zymogen granules, with very few attenu-
ated supporting cells lying between. The histological characters
of the epithelium were not, however, very well preserved in any
of my specimens. The two pouches are closely pressed against
the sides of the cesophagus, and in the species in which they are
longest follow the turns of the latter. Hence, as the esophagus
makes a turn towards the left before it passes down through the
loop of the rectum, the right esophageal pouch generally passes
over to the left side and above the cesophagus, the left pouch
passing to the right and below the esophagus and radular sac.
The cesophageal pouches are relatively short in Alcadia palliata
and of approximately equal width thr oughout, but they are much
longer and diminish in diameter towards their posteri lor extre-
mities in A. hollandi. They are particularly long and of a deep
chocolate colour in Paleohelicina ide ; in most species they are
white in spirit-specimens.

Behind the openings of the cesophageal pouches the cesophagus
narrows somewhat abruptly in diameter; the glandular internal
longitudinal ridges disappear, and are replaced by nine or ten
longitudinal ridges formed by columnar ciliated epithelial cells,
and these may be traced throughout its course to the stomach.
This course is a long one, for the stomach lies aslant on the
lower side of the visceral mass, below the greater part of the
lobes of the liver and below the coils of’ the intestine. As seen
from above and behind, it is a large pyriform sac, the narrower
end lying just above the posterior corner of the pericardium ;
the broader end forming on the right side of the visceral mass a
large rounded prominence which fits into the concavity on the
ventral side of the right columellar muscle. The cesophagus
enters the stomach on the upper side of its broader end, and its
course in the several species examined will be best understood by
reference to Pl. XXXII. figs. 10 to 15. Passing to the left as it
enters the visceral mass, the cesophagus lies above the first coils
of the rectum, then passes below the recurrent coil of the rectum,
and so arriving at the dorsal surface of the stomach runs along
the latter as a flattened tube and opens, as stated above, into
its broader end, dilating considerably just at its point of entrance.
This dilatation, which might almost be described as a diver-
ticulum of the stomach itself, receives right and left the wide
ducts of the liver (Pl. XXXT. fig. 7,and Pl. XXXIT. fig. 16, li.d.).

The internal structure of the eae is very complicated, It
attracted the attention of Isenkrahe, who gave a fairly accurate
description of it. Fig. 7 is a representation of a dissection of the
stomach made from behind and below the visceral mass. The
cesophagus is shown at &., and its entrance into the stomach is
indicated by the arrow. The entrance of the left liver-duct is
shown at li.d.; that of the right liver-duct lies on the far side
ef the prominent curved ridge guarding the entrance to the

MORPHOLOGY OF THE HELICINIDZ. 771

cesophagus. Both the esophageal aperture and those of the liver-
ducts are surrounded by complicated epithelial ridges, which, as
shown in the drawing, converge towards and pass into a deep
groove running along the dorsal side of the narrower pyloric
moiety of the stomach. The edges of this groove are bounded by
two preminent folds; that on the right side (the left in the
drawing) being continued towards the esophageal opening as a
projecting ridge, ending at the side of the aperture in a very
prominent crescentic projection covered by a thick iridescent
cuticle.

The internal surface of the stomach is lined by a mixed glan-
dular and ciliated columnar epithelium, the characters of which
are shown in P]. XX XI. fig.8; the glandular elements predominate
in the esophageal, the ciliated elements in the pyloric, moiety of
the stomach. ‘The epithelial cells are lower in the furrows, taller
and more slender in the ridges, these latter structures being
formed entirely by thickenings of the epithelium, and not by
foldings of the wall of the stomach. The crescentic projection to
the right of the esophageal opening is formed by a local modifi-
eation of the epithelium, the cells of which are here extraordi-
narily long, with nuclei placed about the middle of their length,
and with apparently homogeneous transparent cytoplasmic contents
(Pl. XXXI. fig. 9); they are all of one kind, without any admix-
ture of glandular cells, and, so far as one can judge, they are not
themselves glandular. The free ends of these cells are covered bya
very thick and tough cuticular coat, which stains deeply in hema-
toxylin, brazilin, and other ordinary dyes. The whole structure
corresponds to the “ fiéche tricuspide,” of which the characters
have been thoroughly described for Lamellibranch stomachs,
and which has been noted as occurring in several Gastropod
‘stomachs. There is no definite crystalline style in Helicinide,
but in several specimens that I dissected I found the cavity of
the stomach filled by a semitransparent gelatinous mass, which
appeared to be similar in origin and composition to a crystalline
style. In several specimens I found that the intestinal end of
the groove of the pyloric moiety of the stomach was occupied by
closely compacted fecal pellets or rods, while the cavity below
was filled either by the gelatinous mass above mentioned or by 4
loose mass of semi-digested food. It may be inferred from this
that digestion is effected in the general cavity of the stomach,
and that the indigestible materials of the food are collected into
the dorsal groove and passed into the intestine.

Morphologically, the stomach of the Helicinide closely resembles
that of the Neritide, and further bears a resemblance to the
stomachs of the Fissurellide# and Scissurellide#, which, as in this
case, have a groove leading from the hepatic ducts towards the
intestinal end of the stomach, but no spiral cecum.

The small intestine is a comparatively narrow tube, which after
leaving the pyloric end of the stomach runs back for a short
distance over the dorsal surface of the latter, parallel with the

172 PROF, G. C. BOURNE ON THE

cesophagus, and then turning upwards and forwards describes one
or more convolutions before it passes into the large intestine.
The small intestine can always be distinguished by its white
colour and narrow diameter; it varies considerably in length in
different species, being longest in Alcadia palliata and Orobophana
ponsonbyi, shortest in Hutrochatella pulchella. Internally it is
lined by a columnar ciliated epithelium containing a few gland-
cells, and its internal surface is increased by a well-marked
internal ridge or typhlosole formed by long ciliated cells. The
small intestine passes abruptly into the large intestine, the latter
being of much larger diameter and having pigmented walls. In
all species of Alcadia and Helicina examined, the large intestine
runs forward and to the left below the cesophagus, then bends
abruptly back, forms a wide circular loop which passes above the
cesophagus, in front of the anterior end of the stomach and along
the inner wall of the anterior end of the pericardial cavity: it then
turns downwards and to the right, passes right round the lower side
of the stomach, and mounting upwards again behind it runs in the
right side of the roof of the mantle-cavity to open by the anus,
opposite the right tentacle. In the first part of its course, 7. e. in
the short length between the small intestine and the recurrent
circular loop, the large intestine is lined by very evenly disposed
columnar ciliated cells, among which I could not detect any
gland-cells, and this part of the intestine is not, asa rule, full
of fecal matter. This section of the intestine may be described
as the large intestine proper, to distinguish it from the rectum,
into which, however, it passes without any obvious line of
demarcation. ‘The rectum is always full of feecal débris, and its
epithelium consists of (1) columnar ciliated cells; (2) goblet-
shaped gland-cells filled with coarse granules which stain deeply
in hematoxylin and are therefore probably mucinogenous ;
(3) smaller gland-cells containing small yellow granules. There
is no typhlosole either in the large intestine or the rectum.
Noticing that the coils of the gut differed in the different
species, I have been at some pains to work out this character in
detail, and figs. 10 to 15 (Pl. XXXII.) show the coils charac-
teristic of six different species. An examination of the figures
will give a better idea of the differences than any description.
It will be noticed that there are three main types. In Alcadia
palliata, A. hollandi, Orobophana ponsonbyi, and Paleohelicina
the intestinal coils, though differing in detail, are alike in this
respect, that the small intestine runs back more or less parallel
to the cesophagus over the dorsal surface of the stomach, and
the large intestine makes a bend to the right below the cesophagus
and then, turning back on itself, makes a second bend to the
right above the esophagus. The second type of arrangement
is shown in Lucidella awreola (fig. 14). In this species the
esophagus is not pressed against the dorsal surface of the
stomach, but runs obliquely down to it; the small intestine
passes forward from the pyloric end of the stomach, crosses over

MORPHOLOGY OF THE HELICINIDA. 773

the cesophagus, and curves round it till it nearly touches the
pyloric end of the stomach again; here it passes into the large
intestine, which turns sharply back, passes round the cesophagus
again, and coming to the surface sweeps round to the left in front
of the anterior corner of the pericardium to form the descending
loop of the rectum. This type is easily derived from the first by
the shortening of both the small and the large intestine, in
consequence of which the former is hooked round the cesophagus
and one of the bends characteristic of the first type is suppressed.
The third type, seen in Hutrochatella pulchella (fig. 15), differs
considerably from the other two. The small intestine is even
shorter than in Zuecidella, and the loop formed by the large
intestine and the first section of the rectum lies wholly on the
dorsal side of the esophagus; this condition is clearly due to
the gut being much shorter than in the other types, and it
appears to be quite a constant feature in Hutrochatella. The
intestinal coils of Aphanoconia andamanica ave singularly like
those of Hutrochatella.

I may appear to be giving an undue amount of attention
to characters of no obvious morphological or physiological
importance, but it is just because they may be claimed to be
of importance in the economy of the species that I have spent a
considerable amount of time in working out these details. Hach
species seems to have a characteristic arrangement of the coils
of the intestine, and the arrangement is remarkably constant in
individuals of the same species, allowance being made for
displacements due to the greater or less state of contraction of
the specimens. Closely allied species, such as Alcadia palliata
and A. hollandi, have a very similar arrangement, yet sufficiently
different to allow one to recognize them at a glance after
obtaining some familiarity with their anatomy. Lucidella and
Eutrochatella, both separated from Aleadia and from one another
by distinctive characters of shell, operculum, and radula, differ
in a nearly corresponding degree in the coils of the intestine.
Orobophana and Palcohelicina are Pacific forms which must
have been derived from American Helicine, the latter being
closely related to Alcadia, and they resemble the last named in
the coils of the intestine. To this extent it may be claimed that
two, three, or more characters vary together in these genera ; but
Aphanoconia presents a difficulty, for this genus is far removed
from Hutrochatella in shell and radular characters, and is in these
respects closely related to Paleohelicina, yet its intestine is as
nearly as may be that of an Hutrochatella. As the two genera
cannot possibly stand in close genetic relationship to one another,
the similarity in the pattern of the intestinal coils must be due
to parallelism, similar causes producing similar deviations from
type in the two organisms. It has been shown that the
differences in pattern are attributable to differences in the
length of the large and small intestines, and this is probably
connected with different forms of food. As we are ignorant of

774 PROF. G. C. BOURNE ON THE

the habits of the various species of Helicinide, and do not even
know for certain whether each or any species is restricted to a
particular kind of food, it would be rash to speculate on this
question, but such evidence as I have collected does seem to show
that such apparently trivial characters as the coils of the intestine
are of some physiological importance, and are therefore subject to
the action of natural selection.

The Celom.

The ewlom is represented by the pericardial cavity, which,
although it is of much smaller extent and less complicated than in
the Neritide, is nevertheless a cavity of relatively considerable size,
as may be seen by an inspection of figs. 17 to 20 (PI. XX XIIT.).
As is shown in fig. 1 (Pl. XXX.) it comes close to the surface of
the left side of the visceral mass, and extends forwards nearly
as far as the posterior end of the left columellar muscle and
backwards round the lower side of the visceral mass as far as the
end of the mantle-cavity. It is bounded externally by the very
thin body-wall, posteriorly by the inner wall of the mantle-
cavity, internally by the kidney (figs. 17 to 20). At about the
middle of its length it is a cavity of considerable depth, extending
some way into the visceral mass below the pyloric division of
the stomach. The reno-pericardial canal, which will be described
in connection with the kidney, opens into its right posterior wall,
at some little distance from its hindermost end (fig. 19); other-
wise it is a closed sac containing the auricle and ventricle of the
heart, and does not require further description.

The Hemocele, Circulatory and Respiratory Systems.

The blood-vascular system, as in all Molluscs, consists partly
of large lacunar spaces, which collectively are known as hemocele,
and partly of vessels with definite walls. The hemoccelic spaces
surround the viscera, and there is a specially large lacunar space
below the buccal bulb, in which lie the pleuro-pedal ganglia.
In the visceral mass and in the dorsal region of the body the
hemoceele is largely filled up by the peculiar form of connective
tissue which I have previously described (2, p. 861) as metabolic
tissue. In the Helicinide the tissue is of precisely the same
nature as in Neritidz, and it is not necessary to describe it again.
It evidently consists in large part of reserve tissue, for it is most
abundant in immature specimens in which the gonads and
gonaducts are but slightly developed, and is much less abundant
in sexually mature specimens. This metabolic tissue is specially
concentrated round the larger blood-vessels.

It is not possible to trace the course of the blood-vessels by
dissection of spirit-preserved specimens and only the larger vessels
can be traced in sections. The following account of the
circulation embodies as much information as I have been able to

MORPHOLOGY OF THE HELICINIDA. 775

obtain by reconstruction of sections. The ventricle is continued
forward into a short and wide aorta, which immediately after
passing through the pericardial wall—which it does at about the
level of the hinder end of the left columellar muscle—divides
into three principal branches. The one passes to the right
towards the stomach, and, entering the visceral mass, divides
into a number of branches which are distributed to the stomach,
the intestine, the right lobe of the liver, the gonad, and the
hypobranchial gland and gonaducts. The left branch runs
forward for a short distance and then turns downward into the
visceral mass and chiefly supplies the left lobe of the liver.
A branch is directed towards the posterior part of the left
columellar muscle. The third vessel is an almost direct forward
continuation of the aorta and runs up in the dorsal region of the
body towards the esophagus; it passes above this organ and runs
over the surface of the radular sac, to which organ it becomes
firmly attached at about the level of the hind end of the
pharyngeal bulb. Here it passes into a number of lacunar
passages, supplying the pharyngeal bulb and the cesophageal
pouches, and eventually makes communication with the large
blood-space surrounding the nerve-centres of the head. This
space in turn communicates freely with the lacunz surrounding
the pedal nerve-chords.

In this labyrinth of blood-channels I have not been able to
recognize those by which the blood is collected and brought back
from the various organs to the organ of respiration, the mantle.
The principal hemoceelic spaces or blood-sinuses are the
following: (1) a pedal sinus, surrounding the pedal nerve-cords ;
(2) a subcesophageal sinus, underlying the buccal mass and
esophagus ; this is continued back into (3) a cireum-intestinal
sinus, in which lie the coils of the intestine. and the posterior
part of the radular sac; (4) a peri-gastric sinus surrounding the
stomach ; (5) a recto-genital sinus, running the whole length of
the rectum and gonaduct. The last named is evidently the
pulmonary vein of Isenkrahe (“lings des Darmes zieht sich die
Lungenvene hin”), but it does not carry back blood from the
mantle to the auricle. On the contrary, it is easy to see that
blood passes from it to the numerous fine blood-vessels or
rather blood-spaces, for they have no definite walls, in the roof of
the pulmonary chamber. The efferent pallial vein that collects
blood from the mantle and returns it to the heart is on the opposite
or left side of the mantle-cavity. It isa direct continuation of the
auricle and can easily be traced forward in the left corner of
the mantle-cavity, running along the upper border of the left
columellar muscle (Pl. XXXII. fig. 16, v.pal.) nearly as far as
the thick muscular anterior border of the mantle. It receives
numerous vessels from the mantle, especially in the anterior
part of its course. The blood from the intestinal and perigastric
sinuses does not pass straight to the mantle, but is collected into a
large sinus provided with definite walls (Pls. XX XIII. & XXXIV,

776 PROF, G, C. BOURNE ON THE

figs. 19 & 21, v.ven.), whence it passes by an afferent renal vessel
to the glandular portion of the kidney. So far as I can ascertain,
the blood is returned from the kidney to numerous small vessels
running in the floor of the posterior half of the mantle-cavity,
and is conveyed from these to the auricle by a distinct vein
(Pl. XXXIITI. fig. 20, v.post.) which opens into the hinder part of
the auricle, and receives in addition blood from the roof of the
extreme hind end of the roof of the mantle-cavity. There are
thus two distinct vessels opening into the auricle, the foremost of
which brings back blood from the roof of the greater part of the
pulmonary chamber ; the hindmost brings blood that has passed
through the kidneys, then through the vessels on the floor of the
pulmonary chamber, and in addition a small quantity of blood
from the roof of the extreme hind end of the pulmonary chamber.

The Hxcretory Organs.

The topographical relations of the kidney, as compared with
that of the Neritide, have already been explained (p. 765). Re-
garded in detail, the kidney consists of a thiek-walled glandular
portion and a thin-walled non-glandular portion which serves as a
bladder and urinary duct. The glandular portion is a large and,
roughly speaking, quadrangular sac lying in the lower part of the
visceral mass, below the stomach but above and somewhat to the
left of the lower loop of the rectum. Its posterior and left wall
fits closely against the pericardium and forms the inner boundary
of the latter. The two ends of the sac are produced into large
pockets or recesses, which partly extend round and embrace the
walls of the pericardium (Pl. XX XIII. figs. 18 & 19), and partly
extend upwards round the sides of the pyloric division of the
stomach (Pls. XX XJIL. & XXXIV. figs. 17 & 24). The cavity of
the sac is spacious, and only partially subdivided by folds project-
ing inwards from the wall on the pericardial side; the opposite
wall is not folded. The renal blood-vessels run in these folds.

The whole cavity, including the folds, is lined by a uniform
glandular epithelium consisting of large irregularly shaped cells,
of varying length, their free ends rounded or club-shaped and
often projecting far into the lumen of the sac.

The characters of these cells are shown for Alcadia in fig. 22
and for Lucidella in fig. 23 (Pl. XXXIV.). In all the other
species that I have examined the kidney-epithelinm resembles
that of Lucidella ; it is only in Alcadia that the cells are as long,
irregular, and ameeboid-looking as those drawn in fig. 22. In
both cases the cytoplasm is clear and distinctly and coarsely
vacuolated ; the nucleus spherical, vesicular, with a few granules
of chromatin. The ureter or non-glandular part of the kidney
arises from the upper corner of the left-hand recess of the
glandular sac. Its walls are composed throughout of a non-
ciliated, very low, cubical epithelium, the cells of which are so
much flattened that they might almost be called a pavement-

MORPHOLOGY OF THE HELICINIDA. Ce

epithelium. The ureter is a widish tube which after leaving the
glandular sac turns back to run round the hinder wall of the
pericardium, interposing itself between it and the lower surface
of the visceral mass. After passing from the left to the right
side if mounts upwards again, passes under the reno-pericardial
canal, and opens into the right-hand side of the mantle-cavity
by a thick-lipped slit-like uropore. Asis shown in fig. 23, the
mantle-epithelium is invaginated at the lips of the uropore, and
this invaginated portion is ciliated, but there is no uropore-sac
such as I have described in the Neritide.

The reno-pericardial canal (figs. 19 & 22) opens out of the
lower part of the right-hand recess of the glandular part of the
kidney, and runs straight into the right side of the pericardium
opposite the middle of the expanded base of the auricle. The
canal is short, straight, and narrow, lined by a cubical ciliated
epithelium, the component cells of which are small and bear no
resemblance to the very large ciliated cells lining the long twisted
reno-pericardial canal of the Neritide. The cilia are fine and
directed towards the kidney. A thickening of the epithelium at
the pericardial opening of the canal is suggestive of the presence
of a pericardial funnel. The structure and relations of the kidney
and the reno-pericardial duct are remarkably uniform in all the
species of Helicinide that I have examined.

The Generative Organs.

Thiele (10) has shown that the female ducts are monaulic in
Hydrocena cattaroensis, but diaulic in Helicina kubaryi. Before
my memoir on the Neritide was published I had discovered the
diaulic ducts in Aleadia and Hutrochatella, but, as I omitted to
make mention of them in that place, I must yield priority to
Thiele, whose diagrammatic figure (loc. cit. text-fig. 2) gives a
correct representation of the general relations of the various sub-
divisions of the ducts. But it is almost impossible to construct a
life-hke picture of such complicated organs from a study of
sections, and as I have dissected out the gonaducts, both male and
female, in a number of species and have checked my observations
by the study of sections, I may be pardoned for again taking up
the subject and entering into it at some length. From the
analogy of the Neritidz, in which family the gonaducts exhibit a
considerable range of variation, I expected to discover equally
great differences in these organs in the various genera of Heli-
cinide, but have been disappointed. There are differences, it is
true, but they are slight and do not throw much light upon the
systematic affinities of the various genera studied.

The gonads in all Helicinide lie above and to the right side of
the liver. The ovaries are follicular, and the follicles open into a
large thin-walled chamber which in Alcadia and Hutrochatella is
situated on the right side of the visceral mass, just behind the
posterior end of the right columellar muscle and in front of the

778 PROF. G. C. BOURNE ON THE

muscular partition separating the visceral cavity from the complex
glandular mass formed by the hypobranchial gland and gonaducts
(Pl. XXXVI. fig. 32). The ova appear to go through their matu-
ration-phases in this chamber, as no ripe ova are to be seen in the
follicles. In Alcadia and Futrochatella this ovarian chamber is
sac-shaped and on the right side of the body, but in Pacific and
Oriental species, such as Paleohelicina ide, Orobophana ponsonbyt,
and Aphanoconia gouldiana, the ovarian chamber is produced into
a wide tube which stretches transversely across the body and
receives the products of the follicles of a left ovarian lobe, the latter
being a distinct triangular lobe, projecting from the surface of the
visceral mass and packed close under the left columellar muscle.
This lobe is possibly characteristic of Pacific and Oriental species:
it is absent in Alcadia, and scarcely represented in Luérochatella.
My specimens of Zucidella were all male, and therefore I cannot
say whether it occurs or not in this West Indian genus. The
female gonaducts of Alcadia holiandi, which are to all intents
and purposes identical with those of 4. palliata, are depicted in
Pl. XX XV. fig. 25; and figs.30 to 35 (Pls. XXXV. & XXXVI)
represent selected sections from a series passing horizontally
through the genital complex of a female of the same species,
fig. 30 being the uppermost and fig. 35 the lowest of the series.
Dealing first with the macroscopical characters, the following
organs or parts can be distinguished, and as their shape and
relative positions are clearly indicated in fig. 25a detailed de-
scription will be superfluous. (1) The oviduct od. is a very narrow
duct leading from the ovarian chamber to (2) the dilated or V-
shaped portion of the oviduct, in which a descending limb (od.')
and an ascending limb (od.*) can be recognized. In <dAlcadia
the ascending limb is as long and of approximately the same
diameter as the descending limb: in its lowest third it receives
the short and narrow duct of a globular receptaculum seminis.
(3) The ootype, oo¢., is a long, more or less dilated glandular duct
running parallel with the rectum in the right hand of the roof of
the mantle-cavity, and opening into the latter on the right side by
a relatively narrow aperture ona papilla which lies in a sort of
shallow cloaca formed by the expanded lips of the rectal opening.
Into the posterior end of the ootype open: the ascending limb of
the V-shaped part of the oviduct ; the caecum of the ootype, a sac-
shaped structure of considerable relative size, which lies parallel
to and to the inner side of the V-shaped duct ; thirdly, the short
and rather narrow duct of (4) the vagina, vag. The last named
is a slender duct with thin walls, opening into the mantle-cavity
close to the aperture of the hypobranchial gland. It is continued
posteriorly into a sac, which runs back on the outside of the
V-shaped duct and ends blindly (this sac has been displaced in
fig. 25 to show it more clearly, in its natural position it would be
concealed by the V-shaped duct). It may be called the vaginal
sac. Comparing these ducts with those of the Neritide, parti-
cularly with Paranerita (2, fig. 60), it is clear that the ootype is

MORPHOLOGY OF THE HELICINID. 779

homologous in the two forms, as is also the vagina. The vaginal
sac of Alcadia corresponds to the spermatophore sac of Paranerita,
but in the former genus the duct connecting the vagina with the
ootype has been shortened to such an extent that it is merely a
passage between the two. The V-shaped duct of Alcadia is only
a modification of the oviduct and has no exact homologue in
Paranerita; the position of the receptaculum seminis is also
somewhat different. There is no trace of an oviduco-celomic
funnel in any Helicinid, nor is there any representative of the
crystal-sac. On the other hand, the cecum of the ootype is not
represented in the Neritide, for it would be straining homology
too far to suggest that it is the equivalent of the lower dilated
part of the ductus enigmaticus. But if the morphological com-
parison between the female gonaducts of the Neritide and
Helicinide is fairly obvious, a physiological comparison is by no
means so clear. In the specimen of 4. fellandi whose ducts are
drawn in fig. 25 the hinder moiety of the ootype was filled and
greatly distended bya large mass of spermatozoa held together by
a coagulable substance, and, as shown in the figure, a string of the
same mass of spermatozoa and coagulum extended into and filled
the cecum of the ootype. The vaginal sac was empty, but the
receptaculum was full of spermatozoa. This suggests that the
“‘ootype ” is the copulatory canal or functional vagina, and that
the “‘vagina” may serve for the passage of the ova into the
mantle-cavity. But the probability of such a conclusion is
lessened by the fact that in Aphanoconia merguiensis and 4d.
gouldiana I found the vagina and vaginal sac full of spermatozoa.
[have not found ova either in the oviduct or in the ootype of any
of my specimens; and apparently in the Helicinide fertilization is
not effected by means of spermatophores, for I have found no
trace of such structures. The mass of spermatozoa and coagulum
in Alcadia hollandi cannot be called a spermatophore. The
evidence as to the function of the several parts being slender, and
what there is conflicting, I offer no definite theory on the matter,
but may add that the “ vagina” is evidently a distensible duct, as
it has thin walls, with a very feeble coat of circular muscle-fibres,
but with a number of muscular slips passing from its walls to be
attached to adjacent organs. On the other hand, the ootype has
a thick muscular coat, especially in its hinder moiety, and the
contraction of this muscular coat would expel any contained
material through the external aperture.

A comparison of the gonaducts of the several genera shows
certain differences in detail. The female gonaducts of Hutro-
chatella, a West Indian genus, are shown in Pl. XXXYV. fig. 26.
Their arrangement 1s clearly very similar to that of Aleadia. The
ascending limb of the V-shaped duct is shorter in Lutrochatella
and has more the appearance of a direct posterior continuation of
the ootype. The receptaculum seminis is small and ovoid, but in
the same position as in Alcadia. The vagina is short ; the vaginal
sac of moderate length. The cecum of the ootype is a large

780 PROF. G. C. BOURNE ON THE

flattened sac and extends some way behind the posterior end of
the V-shaped duct.

Of the Pacific and Oriental genera, Aphanoconia (Pl. XXXYV.
fig. 27) most nearly resembles Alcadia and Hutrochatella. 'The
descending limb of the V-shaped duct is large ; the ascending limb
short and scarcely differentiated from the hinder end of the
ootype; it bears a relatively large globular receptaculum seminis.
The vagina is fairly long, and opens by a very short transverse
duct rather high up into the ootype. The vaginal sac is short.
There is no cecum to the ootype. The female ducts are very
similar in all the four species of Aphanoconia that I have examined,
differing chiefly in the relative length of the vaginal sac and
the position of the vagino-ootypal connection.

On the other hand, Palwohelicina (fig. 28) and Orobophana
(fig. 29), while resembling one another, differ in some important
respects from the West Indian genera and from dAphanoconia.
In Paleohelicina ide the ascending limb of the V-shaped duct is
wide and scarcely differentiated from the hinder end of the ootype.
There is no receptaculum seminis, but the ootypal cecum is large,
bilobed at its extremity, lined by an epithelium of peculiar cha-
racter, and filled with spermatozoa. It evidently functions as a
receptaculum seminis. The vagina and vaginal sac are normal.
In Orobophana ponsonbyi the descending limb of the V-shaped
tube is unusually long ; the ascending limb very narrow and short,
and does not bear a receptaculum seminis. The cecum of the
ootype is large, bilobed, full of spermatozoa, and in every respect
similar to that of Palwohelicina. The vagina is long; the vaginal
sac leaves it about halfway between its external aperture and its
connection with the ootype.

I am unable to give a description of the female organs of
Lucidella aureola, as all my specimens were males.

Summing up the above facts, we see that as regards the
structure of the gonaducts there are two types in the genera dealt
with. In the first type there is a receptaculum seminis on the
ascending limb of the V-shaped duct. This type is divisible into
two sub-types: the one, found in Alcadia and Hutrochatella, is
characterized by the large cacal appendage of the ootype; the
other, found in Aphanoconia, has no such cecum. In the second
type, represented in Palwohelicina and Orobophana, there is no
receptaculum seminis on the ascending limb of the V-shaped duct,
but this organ is replaced functionally by the modified bilobed
cecum of the ootype. According to Wagner (12) Palcoheli-
cina stands nearest to Helicina sensu restricto, and therefore
nearer to Alcadia than other Oriental and Pacific forms, but in
the structure of the female gonaducts, dphanoconia stands nearest
to Alcadia, but Paleohelicina with Orobophana stand somewhat
apart.

"The gonaducts of the Helicinide evidently undergo great histo-
logical changes at the onset of sexual maturity. In sections of
immature females of Alcadia and Hutrochatella the ootype is a

MORPHOLOGY OF THE HELICINID®, 781

relatively narrow tube lined by a columnar epithelium, in which
glandular elements are hardly recognizable, and there is a similar
lack of differentiation of glandular cells in the V-shaped duct. On
the approach of sexual maturity the epithelium of the ootype is
enormously thickened, is rich in glandular elements, and is thrown
into complicated folds; at the same time, histological changes
occur in the V-shaped duct. In the breeding-season, when the
ootype is distended as shown in fig. 25 (PI. XX XV.), the epithelial
folds of the ootype disappear, the gland-cells have discharged their
contents, and the epithelium appears shrunken and thinner than
before. In consequence of these changes, it is difficult to give
a consistent account of the histological characters of the ducts :
what follows is based on a series of sections of a mature
specimen of Alcadia hollandi, in which the tissues, thanks to
Mrs. Longstaff’s care, are admirably preserved.

In the distal half of the ootype, that is in the portion marked a
in fig. 25, the epithelium is moderately long and of the mixed
glandular and ciliated kind. The ciliated cells are elongate
columnar, not much attenuated at their bases, their cytoplasm
elear and very finely granular, their nuclei rather large, oval,
situated rather to the basal side of the middle of the length of
the cell. The gland-cells are of nearly the same shape, but of
rather greater diameter than the ciliated cells, their nuclei slightly
larger, situated nearer the bases of the cells, and the cell-body
filled with rather small highly refracting spherules of a greenish-
yellow colour in preparations stained with hematoxylin and eosin.
The hinder moiety of the ootype—namely, that portion filled with
the mass of spermatozoa in fig. 25—shows somewhat different
characters. The supporting epithelial cells are, as before, columnar
and finely granular, but of greater length: they appear to have
lost their cilia over the greater part of the inner wall of the
ootype and to end distally in rounded and somewhat vacuolated ex-
tremities, but as I can find cilia in patches this appearance may be
due to maceration. ‘The gland-cells in this region (P]. XX XVII.
fig. 36) are not very abundant, but characteristic, and presumably
mucinogenous as they stain deeply in hematoxylin. Their basal
ends, resting on the basal membrane, are broad and in each is a
rather small subspherical nucleus, above which the cell tapers to
a fine tube filled with a darkly staining granular material: these
attenuated cell-bodies run between the supporting cells and ter-
minate in swollen extremities filled with coarse deeply staining
spherules. Throughout this region of the ootype the epithelium
is thrown into ridges and furrows, which are partly due to the
folding of the walls, but chiefly to the unequal length of the
epithelial cells. At the hinder end the folds increase and there is
a prominent valve projecting into the lumen and making a com-
plete spiral turn just above the opening of the V-shaped duct.
This opening, guarded by the above-mentioned valve, is narrow.
The upper part of the V-shaped duct is lined by an epithelium of
the same character as that of the hinder moiety of the ootype, the

Proc. Zoot, Soc.—1911, No. LIV. 54

782 PROF. G. C. BOURNE ON THE

supporting cells being here distinctly: ciliated and the deeply
staining mucinogenous cells conspicuous. This epithelium is
thrown into a distinct spiral ridge, which winds round the upper
part of the V-shaped duct nearly as far as the entrance of the
duct of the receptaculum seminis. Here the epithelium changes
its character; the deeply staining mucinogenous cells disappear
and give place to very numerous long tubular gland-cells with large
oval basal nuclei, the tube-shaped cell-body coarsely alveolar and
the alveoli containing large refringent non-staining spherules.
Wedged between these are the ciliated cells, with elongated nuclei
at about the middle of their length, very attenuated basal ends,
and somewhat expanded wedge-shaped distal ends, each with a
distinct striated border anda tuft of fairly long cilia. The recep-
taculum seminis (Pl. XX XVIT. fig. 37) is lined by a low columnar
ciliated epithelium of uniform character. The cell-bodies are finely
granular and stain readily; the nuclei spherical and deeply
staining. Each cell has a distinct striated border and bears a tuft
of long coarse cilia. The spermatozoa in the receptaculum are all
arranged with their heads directed towards the centre, their tails
outwards and entangled among the cilia of the epithelium. The
receptaculum and its duct are invested by a very stout coat of
muscular fibres.

The transition from the upper end of the ascending limb of the
V-shaped duct to the narrow tube of the oviduct is abrupt. The
oviduct is lined throughout by an epithelium consisting of long
columnar cells bearing specially long and coarse cilia. The cha-
racters of these cells are shown in fig. 38 (Pl. XX XVILI_).

The cecum of the ootype is lined by an epithelium differing
from that of the rest of the ootypein the absence of glandular cells.
The walls of the cecum, like those of the hinder moiety of the
ootype itself, are provided with a tolerably thick coat of muscular
fibres, mostly disposed circularly. The muscular coat is not folded,
but the epithelium is disposed in longitudinal ridges due, as seen
in fig. 39 (Pl. XX XVIT.), to the greater length of the cells com-
posing them: this figure is from Orobophana ponsonbyi and not
from Alecadia; in the latter genus the cells are somewhat longer
and more slender, but otherwise similar in character. Asis shown
in the figure, the cells are club-shaped with rounded ends projecting
into the lumen of the cecum; they do not bear cilia. It is the
presence of this characteristic epithelium in the bilobed sac full
of spermatozoa leading into the hinder end of the ootype in Paleo-
helicina and Orobophana which leads me to identify the sac in
question with the cecum of the ootype of Aleadia and Hutrochatella
rather than with the receptaculum seminis, and it is further to be
remarked that in the non-ciliated bilobed sac the spermatozoa are
arranged pell-mell, with their heads and tails in all directions
instead of being definitely oriented as they are in the ciliated
receptaculum seminis.

The distal third of the vagina is lined by an epithelium consisting
for the most part of highly vacuolated clear cells with basal nuclei,

MORPHOLOGY OF THE HELICINIDA. 783

and between these are very attenuated supporting cells. I could
not detect any cilia on the latter.

The proximal two-thirds of the vagina and the vaginal sac
are lined by a non-ciliated epithelium of uniform character, the
details of which were not well preserved in my sections of Alcadia
and Hutrochatella: apparently they had been injuriously affected
by the action of Perenyi’s fluid. In Orobophana ponsonbyi the
epithelial cells of the lower part of the vagina and of the vaginal
sac are squarish in outline, non-ciliated and clearly glandular, for
each contains a number of coarse non-staining spherules. The
cavity of the sac contains a number of spherules of similar
character and among them ropy masses of some coagulable sub-
stance staining faintly in hematoxylin.

In Orobophana the epithelial lining of the ootype differs in some
respects from that described for Alcadia. The supporting cells are
attenuated towards their bases, have long compressed nuclei
about the middle of their length, and are distinctly ciliated. The
gland-cells of the distal part of the ootype resemble those of
Alcadia, but are apparently differentiated to some extent, for the
granular contents of those on the inner side of the ootype nearest
the mantle-cavity are eosinophilous, those on the outer side are
not. Just above the entrance of the vaginal duct the eosinophilous
cells are replaced by mucinogenous cells staining deeply in
hematoxylin. A large spiral flap or valve separates the opening of
the cecum from that of the ascending limb of the V-shaped duct.
The last named is very narrow and invaginated for some distance
into the terminal part of the ootype: it has no spiral epithelial fold,
such as is seen in Alcadia. The gland-cells of the distal limb of
the V-shaped tube are highly eosinophilous.

Thiele (10) has shown that the female ducts of Hydrocena are
monaulic. The external aperture leads into a thick-walled glan-
dular duct, which is clearly homologous with the ootype of the
Helicinide. The lumen of this duct is continued posteriorly into
a fairly wide canal lined by a columnar epithelium devoid of
glandular cells, and this ends in a saccular dilatation of consider-
able size, which is apparently glandular; “sein Epithel enthilt
Kliimpchen von Kornchen.” Thiele identifies this sac and its
duct with the vaginal sac and vagina of Helicina, and regards it
as the homologue of the right kidney of the Trochid, which in
this case has not acquired an independent opening into the mantle-
cavity. In addition to this sac, three other. structures open into
the hinder end of the “ootype” in Hydrocena: on the right a
thick-walled glandular cecum; on the left the oviduct; and
between the two and dorsad of the “right kidney sac” a tubular
receptaculum seminis. Thiele’s homologies seem to be perfectly
just, and after his discovery of the conditions obtaining in Hydro-
cena, | must agree with him in regarding the vaginal sac of the
Helicinidee and the spermatophore sac of the Neritide as the
representatives of the right kidney. But I still beg leave to
differ from his interpretation of the vaginal aperture as the

54*

LA
784 PROF. G. C. BOURNE ON THE

primitive aperture of the right kidney, and of the external aperture
of the ootype as a secondarily acquired separate genital duct. The
conditions in Hydrocena appear to me to be an ample justification
of the argument put forward in p. 873 of my memoir on the
Neritide. Hydrocena is in many respects more primitive and
therefore more nearly related to the ancestral Neritoid stock than
the Helicinide, and Thiele himself points tothe generative organs
as one of the evidences of primitive organization.

In his memoir entitled “« Die systematische Stellung der Soleno-
gastren und die Phylogenie der Mollusken” (9) Thiele, after de-
scribing the male organs of Septaria, gives the following account
of the male organs of Helicina (= Waldemaria Wagner) japonica :
“ Bei Helicina ist der miinnliche Geschlechtsapparat merklich
einfacher, der Samengang ist nur sehr wenig aufgekniuelt und
weiter, er mundet in den Driisengang von unter und rechts nicht
weit vor seinen Hinterende. Der letztere ist bedeutend einfacher
als bei Vavicella, durch Falten streckenweise etwas zertheilt,
doch scheint das driisige Epithel trotz geringer Verschiedenheiten
an manchen Stellen in Wesentlichen gleichartig zu sein. Die
Driisenzellen liegen durchweg zwischen den Stutzzellen. - Dieser
Gang reicht weit nach vorn in der Mantelhohle doch habe ich ein
besonderes Kopulationsorgane nicht wahrgenommen. Die ekto-
dermale Driise, welche rechts von Hinterende des Driisenganges
(Prostata) in die Mantelhéhle ausmiindet, erstreckt sich, in dem
sich allmihlich grosser wird, weit nach hinten, wo sie neben dem
Hinterende der Niere aufhért. Nach ihrem Bau ist an ihrer
ektodermalen Herkunft nicht zu zweifeln, da sie zwischen den
grobkornigen Driisenzellen deutliche Stutzzellen enthalt. Sie mag
als eine Art von Manteldriise ihnlich der Hypobranchialdriise
anzusehen sein: ihre Funktion ist unbekannt.”

The ectodermal gland referred to in this passage I have already
described as the hypobranchial gland, fully agreeing with Thiele’s
interpretation of it. It has no connection, with the generative
organs. In the six genera that I have examined the male ducts
are, with the exceptions to be mentioned hereafter, very much
alike, but not quite so simple as Thiele’s description would lead
one to suppose, and presumably Waldemaria japonica has undergone
some simplification in these organs, for so accurate an observer
cannot have overlooked the accessory organs that I am going to
describe.

The testis, like the ovary, is follicular in structure ; fine thin-
walled ducts converge from the follicles and unite in the mght
side of the visceral mass to form the sperm-duct. The last-named
organ occupies a position similar to that of the oviduct in the
female: in immature specimens it is slender, slightly convoluted,
and lined by a columnar ciliated epithelium. In mature specimens
its middle portion is greatly distended by spermatozoa, is consider-
ably convoluted, and the ciliated epithelial lining is no longer
distinguishable. The sperm-duct tapers somewhat and, as Thiele
clescribes, opens into the lower and right side of a long glandular

MORPHOLOGY OF THE HELICINID. 785

thick-walled sac, some little distance in front of the hinder end of
the latter. This thick-walled sac, which is evidently the repre-
sentative of the ootype of the female and of what I have called the
terminal chamber in the Neritide, runs forward in the roof of the
mantle-cavity, below and to the right side of the rectum, and
opens into the mantle-cavity by a terminal pore situated close to
the anus. At about one-third of its whole length from the
external aperture, the thick-walled sac—which I shall call the
terminal sac of the sperm-duct or, more shortly, the terminal sac
—~is Joined by another sac of considerable diameter. ‘This second
sac, which I shall call the diverticulum, opens into the terminal
sac by a wide aperture, and runs back close to the right side of
the latter, to end blindly, sometimes just in front of the entry of
the sperm-duct into the terminal sac (Aphanoconia gouldiana,
Pl. XXXVIT. fig. 41); in other cases, however, it extends as far
back as the hinder end of the terminal sac, and may even project a
little beyond it (Alcadia hollandi, fig. 40). The anterior third of
the terminal sac, in front of the entry of the diverticulum, exhibits
three or four deep transverse constrictions: in section it is round
or oval, and the lumen is partly occluded by deep longitudinal folds
projecting into it. These folds are covered by a mixed glandular
and ciliated epithelium: the ciliated cells of the familiar kind with
attenuated basal ends, the gland-cells tubular with basal nuclei and
vacuolated cell-bodies, in which no secretory granules could be
distinguished in well-preserved specimens of Hatrochatella. In its
posterior two-thirds the terminal sac is laterally compressed so as
to be elongate oval in section, and it gives off from each end of the
oval and from the adrectal side numerous short hollow cecal out-
growths which are sometimes branched, especially in Aphanoconia
(fig. 41). Internally the longitudinal epithelial ridges die out in
the posterior two-thirds of the terminal sac, but the epithelial
lining both of the cavity of the sac and of the cecal outgrowths
is of very nearly the same character as that of the anterior third.
The supporting cells are distinctly ciliated. The epithelium of the
diverticulum difters from that of the terminal sac only in the fact
that the gland-cells are full of eosinophilous granules, and the
cilia of the supporting cells are longer and rather coarser. The
above characters hold good for all the species that I have examined,
the differences between them being too slight to deserve mention.
It may be noted that in Alcadia the hinder moiety of the diverti-
culum is constricted at very regular intervals (Pl. XXX VIT. fig. 40).
Both in Hutrochatella and Lucidella I have found in sections a
second diverticulum in the form of a slender thin-walled tube
opening into a recess of the terminal sac at the same level as, but
on the opposite side to, the diverticulum above described. The
walls of the recess are lined by a glandular epithelium staining
deeply in eosin. The narrow tube runs back in the mantle-wall
nearly parallel to the terminal sac and ends blindly just in front
of the aperture of the hypobranchial gland. Its hinder end
touches and appears to be adherent to the sub-epithelial muscular

786 PROF. G. C. BOURNE ON THE

wall of the mantle-cavity, but there is no aperture into the mantle.
I have not been able to find this tube in other species, but this
may be due to the imperfection of my sections and to the fact that
it is too small to be recognizable in dissections. It is a well-defined
structure in Hutrochatella and is lined throughout. by a non-glan-
dular cubical epithelium. I am inclined to the opinion that the
anterior third of the terminal sac of the male is the equivalent
of the body of the ootype of the female; the diverticulum of the
male represents the cecum of the ootype, and the narrow tube
(Pl. XXX VIT. fig. 42, 4.7.) represents the vagina of the female,
but has lost its opening into the mantle-cavity. If this iden-
tification is correct, a relic of the right kidney-sac is retained in
the male, at least in Hutrochatella pulchella and Lucidella aureola.

It may be noted that Isenkrahe’s drawing of the male organs
of Helicina titanica is very nearly correct.

The Nervous System.

Nobody has given a detailed account of the nervous system of
any Helicinid since Bouvier dealt with this subject in his
classical memoir on the nervous system of Prosobranch Gastropods
(3). In that work he gives an elaborate figure of the nerve-
centres and principal nerve-trunks of Helicina sagraiana dOrb.,
and also several figures of the buccal ganglia of the same species
and of the cerebral and pleuro-pedal centres of H. brasiliensis
Gray. As is always the case, Bouvier’s figures possess a high
degree of accuracy, and if I have some criticisms to offer, they
must not be taken as depreviatory of his excellent work, but
as an elaboration of it, rendered possible by careful study of
sections and by the opportunities for exceedingly fine dissection
afforded by the Braus-Driiner microscope.

In the first place, it was necessary to determine whether the
supra-intestinal nerve exists in the Helicinide. Bouvier had
failed to find it in the Neritide, and when in a subsequent memoir
he announced its discovery in the latter group, he hazarded the
opinion that it would probably be found in the Helicinide.
But it does not exist in these pulmonate rhipidoglossates ; it has
‘disappeared in them as completely as the organs with which,
when present, it is associated, the ctenidium and the osphradium.
I can speak with certainty on this point, for I have made so
many dissections and have studied such a sufficient number of
serial sections that I could not have overlooked it if it were
present.

In the second place, I am unable to verify some of the details of
Bouvier’s figure of the nervous system of 7. sagraiana. In none of
the species that I have studied are the pedal, pleural, and subintes-
tinal centres as distinct as shown by him. As may be seen in
fig. 44 (Pl. XX XIX.), the pleural ganglia are ill-defined swellings,
scarcely distinguishable from the swollen anterior ends of the
pedal cords, and the subintestinal ganglion is so intimately fused

MORPHOLOGY OF THE HELICINID®. 787

with the pleurals that it is unrecognizable as a separate ganglion,
even in sections (Pl. XX XVIII. figs. 45 to 52). The pedal cords,
though not so widely separated in any of the species of Heli-
cinide that I have dissected as in the Neritide, are not so closely
approximated as shown in Bouvier’s figure. They are fairly
close together and nearly parallel to one another in Hutrochatella
pulchella, somewhat more divergent in Alcadia hollandi, further
apart in A. palliata, and widely divergent in <Aphanoconia
andamanica. The actual amount of divergence or approximation
is, however, undoubtedly dependent on the degree of contraction
of the muscular mass of the foot, and is a character cf no great
importance. If the foot is much contracted the pedal cords are
approximated and the numerous and slender pedal commissures
are arched : if the foot is relaxed the cords are turther apart and
the commissures are pulled out straight. I cannot but think
that Bouvier has exaggerated the length of the cerebro-pleural
and cerebro-pedal connectives. In none of the species that I have
examined are they appreciably longer than the antero-posterior
diameter of the cerebral ganglia, and in some species, e. g.
Lucidella aureola, they are very short, but it is, of course, possible
that they are unusually long in H. sagraiana. In respect of the
nerves issuing from the ganglionic mass formed by the fusion of
the anterior ends of the pedal with the pleural centres, Bouvier,
while otherwise exact, makes one important omission. He does
not figure or describe a relatively large pair of nerves which
originate one on each side of the most anterior pedal commissure
from the dorsal surface of the swollen anterior ends of the pedal
cords. Each of these nerves (Pl. XX XIX. figs. 43 & 44, m.op.)
passes outwards and backwards, penetrates the muscular wall of
the body, and passes to the muscles of the operculum, hence the
nerves in question may be called the opercular nerves. That of
the right side breaks up into a number of fine twigs in the
opercular muscles, that of the left side gives off a stout branch
which passes to a peculiar hollow organ connected with a plate of
cartilage near the left corner of the opercular Jobe. This organ
will be described in detail further on.

The otocysts are situated just above the origins of the two
opercular nerves, and are therefore on the dorsal side of the pedal
ganglia, as is shown in fig. 44. Bouvier, describing the otocysts
of Helicina brasiliensis says ‘elles sont situées sous les cordons
pedieux ”: and it is true that in a retracted specimen, in which
the head has been drawn back behind the foot, so that the
pedal cords appear to lie in front of the cerebral ganglia, one
does find the otocysts below the pedal cords, when making a
dissection from the dorsal side. But in such a case the sole of
the foot is uppermost, and the morphologically dorsal side of the
pedal cords is turned downwards, this change of position being
very puzzling to the observer both in sections and dissections.
It is worth remarking in this connection that in my experience
the small Polynesian genera, Aphanoconia, Palcohelicina, and, in

788 PROF, G. C. BOURNE ON THE

a lesser degree, Orobophana, when they withdraw themselves into
their shells, do not contract themselves as much as the West-
Indian species. The sole of the foot in these Polynesian genera
is longer and narrower than in the West-Indian genera, the
columella: muscles longer and inserted further back from the
mouth of the shell. ‘There is therefore more ample room for the
head and foot in the last whorl of the shell, and when the animal
retracts itself the foot is scarcely at all contracted, but simply
slides back with its sole applied to the outer side of the shell till
the operculum borne on the broad opercular lobe closes the
aperture. Specimens of these genera, when extracted from their
shells, do not present the deformed appearance of an Alcadia or
an Hutrochatella, the pedal nerve-cords are not turned forward
with their morphological surfaces reversed, and if the animals
were only a little larger they would be much easier to dissect
than their American congeners. It is possible, and even probable,
that these different modes of retreating into the shell, which are
themselves dependent on the varying length and points of
insertion of the columellar muscles, are correlated with the
different forms of operculum upon which Wagner has founded
his system. At any rate, they are consistent with it, but I have
not been able to follow out this problem in detail.

To return to the nervous system. The opercular nerves must
not be mistaken for the parietal nerves correctly described and
figured by Bouvier and labelled d', e’. The parietal nerves
(Pl. XX XIX, fig. 44, n.par.) are much more slender than the
opercular nerves and originate, as shown in fig. 44, from the
pleural centres, between the great pallio-columellar nerves and
the cerebro-pleural connectives. They pass to the muscular walls
of the head behind the tentacles. From the ventral side of the
swollen anterior ends of the pedal cords, in the same cross-section
as the opercular nerves, a rather stout pair of nerves originates
near the middle line; these nerves, which are shown in section
in fig. 46 (Pl. XX XVIIT.), pass to the pedal gland and appear to
be specially connected with that organ.

As regards the subintestinal nerve and its distribution, I am
unable to bring my observations into agreement with those of
Bouvier. The short nerve connecting the subintestinal with the
left pallio-columellar nerve-trunk does not appear to be a constant
feature. J have found such a connective in a single specimen of
Alcadia palliata, and in that one instance it is much closer to the
pleuro-pedal centres than is shown by Bouvier. But I can find
no trace of it in any other specimen that I have examined. I am
unable to find any trace of the visceral nerve labelled j’ in
Bouvier’s figure, which he describes as given off from the left side
of the subintestinal at some distance from the origin of the latter,
and in general my observations on the subintestinal and visceral
nerves differ so much from his that a detailed account is necessary.
The figure illustrating this account (PI. XX XIX. fig. 43) is founded
upon dissections, and the ultimate ramifications of the principal

MORPHOLOGY OF THE HELICINIDA. 789

nerve-branches have been traced in sections. The subintestinal
trunk in Alcadia palliata, A. hollandi, and Lutrochatella pulchella,
after leaving the subintestinal ganglion, which latter is unrecogniz-
ably fused into the pleural centres, courses along the floor of the
anterior division of the general body- -cavity, below the pharyngeal
bulb, the radular sac, and cesophagus. It keeps closer to the right
than to the left columellar muscle, and on reaching the posterior
end of the muscle it doubles the angle between it and the visceral
mass, and entering the latter turns to the left and enlarges to
form a ganglion of some size from which several nerves are given
off. That this isa true ganglion-centre, and not a mere nodal thick-
ening at a point from which several nerve-branches originate, is
denionstrated by the considerable sheath of nerve- ganglion cells.
The principal nerves issuing from the ganglion, in ade inn to the
main trunk, are the following :—a small nerve, v', which passes to
the right near the surface of the visceral mass and is distributed
to the gonads and liver. Another small nerve, v°, which passes
to the left of the visceral mass and appears to innervate the right
lobe of the liver and-surface of the stomach. A stout nerve,
n.gen., which runs to the right, passes above the oviduct or
sperm-duct, gives off a large branch to the mucous gland, and
turns forward to break up into twigs on the posterior part of the
complex of genital ducts: this is the genital nerve, and it
identifies the ganglion from which it originates as the visceral
ganglion. There is no separate genital ganglion as in_ the
Neritide. The hinder end of the visceral ganglion is continued
into a rather stout nerve, which may be regarded as the
continuation of the main trunk of the subintestinal: this passes
through the liver and helow the small and large intestine, and
turning towards the left it passes towards the lowes border of the
right moiety of the kidney, near which it enlarges to form a small
but distinct ganglion, which I take to be the representative of the
elongated visceral ganglion of the Neritide. From this ganglion
small nerves are given off to the liver and kidney, and a larger
nerve passes below the kidney, skirts the uropore, and can be
traced beyond as far as the auricle of the heart, at which point it
ceases to be recognizable. It is a matter of extreme difficulty to
follow the above-mentioned nerves through the liver and intestinal
coils by simple dissection, and I have only been able to make sure
of their ultimate course by the study of serial sections.

After this criticism of the general characters of the nervous
system, | may return to the consideration of some special details.
The pleuro-pedal centres and with them the subintestinal
ganglion are, as I have already said, so intimately fused as to be
practically indistinguishable as separate ganglia. This fusion is
brought out in a striking manner in sections. Figs. A5 to 52
(Pls XXXVIIL) represent selected members of a series of nearly
transverse sections through the pleuro-pedal centres of Alcadia
hollandi. Fig. 45 represents a section through the pedal cords at
the point where they begin to diverge from one another. Dorsad

790 PROF. G. C. BOURNE ON THE

of them are the otocysts, o¢.; ventrad of them is the pedal gland.
Each cord consists of a core of nerve-fibres and dendrites surrounded
by a cortical layer of nerve-ganglion cells. The latter send in
lateral horns at about the middle of the outer side of each cord
in such a manner as to divide the central core into upper and
lower moieties, which have been identified by French authors
with the pleural and pedal sections of the cords respectively.
This interpretation, however, does not appear to me to be well
founded. Fig. 46 represents a section somewhat further
forward than that in fig. 45; it passes through the hinder part
of the anterior pedal commissure, and includes the roots of the
two nerves of the pedal gland and of the right opercular nerve.
The nerve-fibres of the former are seen to be supplied from two
areas of the cortical layer lying respectively on the outer and
inner sides of the ventral side of the cord. The opercular nerve
receives its fibres partly from a centre on its own side, partly
from a centre on the opposite side of the cord, the latter fibres
crossing over in the commissure. Below them is a stout band of
commissural fibres connecting the lateral horns of nerve-ganglion
cells with one another. Above the median ventral raphe is a
thick mass of nerve-ganglion cells. In a section somewhat
further forward (fig. 47) the ventral raphe has disappeared and
the mass of ganglion-cells above it is only represented by two
small islets of nerve-ganglion cells, which are separated from the
ventral surface by two well-defined bands of nerve-fibres start-
ing from the ventro-lateral groups of nerve-ganglion cells and
passing towards the centre partly decussate, partly sweep round
the islets to curve round to the lateral horns of the ganglion-
cells. Above these curved bands is the well-defined transverse
commissural band of nerve-fibres, and above this again there is
ow either side a centre, consisting of nerve-fibres overlaid by a
layer of ganglion-cells, which is seen to be connected with the
origins of the opercular nerves. Above the nervous mass a pair
of muscular cords passing from the otocysts towards the centre
should be noticed.

In the next section (fig. 48), taken some little way further
forward, the two little islets of ganglion-cells lying opposite the
lateral horns are still visible. Below them are transverse bands
of commissural fibres. Between them is seen the most anterior
part of the decussating tract observed in the previous section.
After decussation the fibres sweep out right and left to the dorso-
lateral regions. In the mid-dorsal line is a deep and wide groove
into which the muscular cords noted in the last section are
entering. I regard the whole of the sections hitherto described
as belonging to the pedal centres. The next section (fig. 49)
shows that the dorsal groove containing the two muscular cords
has been converted into a canal by the upgrowth and dorsal
union of the nervous tissue. All that lies below this canal
belongs to the pedal centres; all that lies to the sides of and
above the canal belongs to the pleural centres.

MORPHOLOGY OF THE HELICINIDAD. 791

In the pedal centres a prominent bundle of nerve-fibres 1s
being formed on either side of the middle line: these when traced
forward prove to be the origins of the cerebro-pedal connec-
tives. Above them are the remains of the transverse anterior
pedal commissure. Laterally, above the lateral horns of the
ganglion-cells two other tracts of nerve-fibres are making their
appearance: these have evidently received large contributions
from the decussating fibres noticed in the previous sections, and
when traced forward they prove to form part of the cerebro-
pleural connectives. The central canal containing the two muscular
cords is surrounded by a layer of nerve-ganglion cells, thickest
on the ventral side, and above the canal are seen tracts of nerve-

fibres originating from the lateral horns of either side and passing
towards the middle line. Above them, again, is a small transverse
band of commissural fibres, and above these a fairly thick layer
of ganglion-cells, which, as may readily be seen by comparing
this with the preceding sections, is something added to what was
there before, and is, in fact, the layer of pleural ganglion-cells.
The next three figures (50, 51, and 52) explain themselves. In
fig. 50 the pedal centres are diminishing rapidly in volume. On
the left side, which, owing to the sections being somewhat oblique,
is rather behind the right side, a large tract of vertical nerve-
fibres is seen passing from the pedal to the pleural centres. To
the outside of this above the lateral horn are tracts of fibres some
of which run upwards and will pass into the left pallio-columellar
nerve, others will be continued into the right cerebro-pleural con-
nective and the right parietal nerve. On the right side the origins
of the cerebro-pedal and cerebro-pleural connectives are well
defined. Between them lie the ganglion-cells of the lateral horn,
and above the nucleus of the cerebro-pleural connective is a second
lateral ingrowth of nerve-cells, belonging to the pleural ganglion.
The pleural centre of nerve-fibres is well masked, and from it a
stout band of commissural fibres passes above the central canal to
the pleural centre of the other side. This is the pleural com-
missure, which I have already described in Neritide. Fig. 51
shows the cerebro-pedal and cerebro-pleural connectives in cross-
section. The central canal with its contained muscular slips,
having passed through the ring formed by the pleural and pedal
centres, emerges as a groove on the ventral side. The origins of
the right parietal and right and left pallio-columellar nerves are
clearly visible, and it may be seen that both pleural centres are
contributing fibres which, passing to the mid-dorsal line, form
the origin of the subintestinal nerve. It should be observed
that some fibres from the last named pass directly into the roots
of the two pallio-columellar nerves. In fig. 52 the origin of the
subintestinal nerve is distinct, and both it and the origins of
the two pallio-columellar nerves appear to be imbedded in a mass
of ganglion-cells in which the limits of the right and left pleural
and the subintestinal ganglia can be traced with the aid of a
little exercise of the imagination.

(2 PROF. G. CG. BOURNE ON THE

From what precedes, it follows’ that the pleuro-pedal centres are
extremely complicated and are largely composed of definite tracts
of nerve-fibres, some commissural, some decussating, some passing
into the nerves issuing from this region. It is noticeable that there
are several connections of a complex kind between the pleural and
pedal centres as well as between the right and left pleural centres
and the right and left pedal cords, and that there are evidences of
numerous nervous relays throughout the region illustrated. No
doubt, on analysis, these apparently complicated nerve-tracts can
be reduced to five groups: (1) the pedal commissure, containing
both direct and decussating tracts of fibres; (2) the pleuro-pedal
connectives ; (3) the cerebro-pedal and cerebro-pleural connec-
tives ; (4) the pleural commissure, peculiar to the Neritide and
Helicinide ; (5) the subintestinal connectives, derived from
both right and left pleurals. Thus, the paths of the nerve-tracts
might be described as normal, and consistent with our knowledge
of the usual connections between the chief nerve-centres of
Molluscs ; but.a study of sections will show that the arrangements
are not so simple as might be inferred from a superficial exami-
nation of the ganglia and their commissures and connectives.
For example, it appears that the cerebro-pleural connectives
contain fibres derived from the pedal centres and that all the
principal nerves contain fibres derived from two or more areas in
the fused pleural and pedal ganglion-mass. I have been unable
to pursue the subject further at the present time, and it is diffi-
cult to make any further progress because of our ignorance of
the physiology of the molluscan nervous system. ‘The main
nerve-trunks must contain both afferent and efferent nerves, and
it seems evident that these pass to different areas of the cortical
layer of ganglion-cells, but as at present we have no means of
distinguishing between these two kinds of fibres further analysis
of the details of the nervous system is impossible. It may be of
use to future workers on this subject to remark that there are two
kinds of ganglion-cells in the cortical layer: larger cells with
clear nuclei staining faintly in hematoxylin, and much more
numerous smaller cells with deeply staining nuclei. It should be
possible to trace the connection of the nerve-fibres with these
different kinds of cells, but such an investigation demands fresh
material, and could form no part of the present work on the
Helicinidee.

As Bouvier found considerable differences in the size and shape
of the cerebral ganglia in Helicina sagraiana and H. brasiliensis,
I have studied these centres with care in the hope that I might
discover characters of classificatory value, but I have been no
more successful here than I was in the case of the genital ducts.
The characters to which Bouvier draws special attention are the
relative size of the cerebral ganglia (enormous in /. brasiliensis) ;
the size and shape of the labial lobe; the origin of the labio-probos-
cidean nerves, which all spring from the labial lobe in H. brasz-
liensis, but only one has this origin in H/. sagraianu. As regards

MORPHOLOGY OF THE HELICINIDE. 793

the relative size of the cerebral ganglia I find that the proportion
of each cerebral ganglion to the pleuro-pedal mass is expressed by
the following figures :—in Alcadia palliata, 4; Alcadia hollandi,
+33 Hutrochatella pulchella, %; Lucidella aureola, 3; Paleo-
helicina ide, nearly +; Orobophana ponsonbyi, 4; Aphanoconia
gouldiana, +3; Aphanoconia rogersit, 2. The relative length of
the cerebro-pedal and cerebro-pleural connectives varies greatly :
they are longest in the two species of Alcadia and in Paleo-
helicona ide ; of moderate length in Orobophana, Aphanoconia,
and Hutrochatella ; extremely short in Zucidella aureola. I have
counted four labio-proboscidean nerves on each side with more
or less certainty in all the species examined with the excep-
tion of Aphanoconia rogersii, in which there appear to be five.
Of these, counting from above downwards, the first and third
are invariably stout nerves which branch soon after their origin ;
the second and fourth are slender and only divide into branches
at their extremities. The labial lobe in all the species at
my disposal has the form of a rounded boss projecting inwards
from the antero-inferior edge of the ganglion: the shape and
relative size of this lobe differ somewhat in the various species,
but the differences are too slight to express in words. Fig. 53
(Pl. XX XIX.), representing the left cerebral ganglion of Palco-
helicina ide, and fig. 54, representing the same ganglion in Aleadia
palliata, show the extremes of difference in shape observed by me,
and, on the whole, the cerebral ganglia of the Polynesian species
resemble those of Palewohelicina, those of the West-Indian species
those of Aleadia. ‘The cerebral ganglia of the Helicinide, wedged
in as they are between the anterior end of the pharyngeal bulb
and the walls of the head, are nearly flat, the labial lobe projecting
inwards beneath the pharyngeal bulb. Because of their flatness
they are very readily stained and mounted as transparent objects,
and figs. 53 & 54 give some idea of the complexity of the nerve-
tracts and centres within the ganglion. One may distinguish an
ocular centre, which is large relatively to the size of the ocular
nerve, a tentacular centre, small relatively to the size of the
tentacular nerve, and a relatively smaller commissural centre.
Hach of the labio-proboscidean nerves has a more or less well-
defined centre of its own, but the buccal and labial commissures
have no distinct centres at their origin. There is, further, a
median lobe which possibly serves as a relay for various nerve-
tracts running into and around it. ‘It is evident that the nerve-
fibres of the cerebro-pleural connective make direct and intimate
connection with the ocular and tentacular lobes, and that a
stout band of fibres curves round from the root ef the cerebro-
pleural connective to the base of the labial lobe, receiving on its
way an accession of fibres from the cerebro-pedal connective.
This nerve-tract makes connections with the centres of origin of
the buccal and labial commissures and the labio-proboscidean
nerves, and a well-defined curved band of fibres sweeps round
from the origin of the first labio-proboscidean nerve to enter the

794 PROF. G. C. BOURNE ON THE

cerebral commissure, the last named also having connections with
the tentacular and median centres. Evidently, the cerebral
ganglia have undergone a high degree of concentration and
integration, but it is to be remarked that if the labial lobe as
figured for Alcadia were pulled out towards the bottom of the
picture, it would form an elongated labial process from which
the labio-proboscidean nerves would be given off at intervals, as
is the case in Zrochus and Turbo. In other words, the labial
lobe, as it seems to me, is represented, not solely by the little
projecting boss from which the labial commissure originates, but
by all that part of the ganglion that hes below a line drawn from
the lower side of the origin of the cerebral commissure to a point
just above the origin of the buccal commissure.

The buccal ganglia, with their commissures and the nerves
originating from them, have been very correctly figured by
Bouvier.

The organ which I have mentioned above as connected with a
branch of the left opercular nerve is shown in section in fig. 55
(Pl. XL.). It is deeply embedded in the muscular tissue of the
opercular lobe and lies to the left of the anterior end of the
lobe close to the origin of the left columellar muscle. It consists
of a cartilaginous plate of subtriangular form, the edges of the
anterior apex inrolled ventrally and eventually fused so as to form
a short conical tube. This cartilage forms, as it were, the cover
of a flattened sac (fig. 55, sac.) lined throughout by an epithe-
lium which is thin and composed of a single layer of somewhat
flattened cells on the side attached to the cartilaginous plate, but
thick and composed of columnar cells bearing short stiff cilia on
the opposite side. Anteriorly this is continued into the tube
formed by the inrolled edges of the cartilaginous plate, and here
the columnar ciliated cells form a nearly continuous lining to
its cavity. Posteriorly, as shown in fig. 55, the ciliated cells
forming the floor of the sac (the reader must understand that the
figure is reversed, so that the ventral side is uppermost) rest on a
thick basement-membrane, from which a broad band of muscular
fibres spreads to be attached partly to the muscular wall of the
left side of the neck, partly to the bands of muscular fibres passing
over the otocysts. Into this muscular band the large branch of
the left opercular nerve penetrates. Taking an anterior course
this nerve gives off a branch to the opercular muscles, but its
main trunk is directed towards the anterior tubular end of the
organ under consideration, and there passes through a small
perforation in the cartilaginous wall and is distributed to the
ciliated columnar epithelium lining the cavity of the sac. Dorsally
the cartilaginous plate is connected with the left columellar
muscle by a stout muscular band passing obliquely outwards.
Above this band are seen in fig. 55 the sections of two convolu-
tions of a coiled glandular tube, which on the one hand communi-
eates by a very narrow duct with the anterior tubular end of the
above-mentioned sac, and on the other hand opens to the exterior,

MORPHOLOGY OF THE HELICINIDA, 795

near the left anterior edge of the operculum, between it and the
membranous flap that surrounds the opercular lobe. This tube is
lined throughout by a glandular epithelium composed of rather
tall goblet-cells with deeply staining basal nuclei and clear cell-
contents.

If, now, we enquire into the morphological significance of this
peculiar organ, I think there can be little doubt as to its homology.
It occupies the same position and receives the same nerve-supply
as the crypt into which fits the curved process of the operculum in
the Neritide. This process is no longer to be seen on the
operculum of the Helicinide, but from a study of the muscular
attachments in the two groups I am inclined to think that the
cartilaginous plate described above represents its inner extremity,
all direct connection with the operculum being lost. In
connection with it new structures have been formed, viz. the sac
and the glandular tube. What its physiological significance is it
is hard to say. It is clear that, by contraction of the muscular
bands attached to it, the sac may be widely dilated, and when
dilated, air must flow into it through the glandular tube. The
abundant nerve-supply and the character of the columnar epithe-
lium bearing short stiff cilia suggest that the sac has a sensory
function, and it is possible that it may be a special sense-organ,
likely enough of an olfactory character, which enables the animal
to receive impressions from the external world when retracted
into its shell. For the opening of the glandular tube is in such
a position that it would open to the outer air whenever the
opercular plug was ever so little loosened.

The pedal gland, an organ which is absent in the aquatic
Neritidee, is largely developed in the Helicinidee. Its position and
general structure are roughly indicated in fig. 45 (Pl. XX XVIII).
It consists of a main duct below the pleuro-pedal nerve-mass and
extending some little way but not far back below and between the
pedal cords. The duct is lined by a columnar ciliated epithelium
raised on the ventral side into two prominent ridges, one on either
side of the middle line. Surrounding the duct are bunches of
unicellular glands, which penetrate among the interlacing muscle-
fibres of this region of the foot. Kach unicellular gland is
prolonged into a fine duct which passes between the epithelial
cells of the main duct and opens into its lumen. The histology
of this gland is reminiscent of that of the byssus gland of
lamellibranchiate molluscs, described by me in another place.
The main duct opens at the anterior end of the foot, in the mid
line, below the snout.

Mention may be made here of the large mucous gland which I
agree with Thiele in identifying as the hypobranchial gland. It is
of relatively enormous size in the Helicinide, but in structure and
position does not differ much from what I have described for the
Neritide. It opens into the mantle-cavity (Pl. XX XV. figs. 28
& 30) just in front of the aperture of the vaginal duct and, as shown
in figs. 30 to 35, it forms a considerable glandular mass bulging”

796 PROF. G, C. BOURNE ON THE

into the mantle-cavity to the left side of the genital ducts. As
may be seen in figs. 16-20 (Pls. XXXII. & XXXIIT.), it extends
far beyond the hind end of the genital ducts, accompanying the
rectum in its course, and lying to the right side of the kidney
at the hinder extremity of the visceral mass. In the section
depicted in fig. 20, which passes throngh the rectal coil at the
bottom of the hind end of the visceral mass, the hypobranchial
gland on the right side of the rectum appears to form a pair with
the kidney on the left, but there is, of course, no relationship
between the two organs. Throughout its course the hypobran-
chial gland consists of an irregularly folded flattened sac, from
which short glandular diverticula are given off in all directions.
In its terminal part, as is shown in figs, 18-20, the gland appears
to be differentiated into two portions: one lying nearer the
cesophageal end of the stomach is lned by an epithelium loaded
with fine dark granules; the other portion, lying nearer to the
mantle-cavity, is lined by an epithelium of the character shown
in Pl. XX XIX. fig. 56. It is made up of large glandular cells,
oblong in outline, and filled with a highly refracting granular
substance which, when the cells are ready to discharge their
contents, is accumulated into oval pellets, as shown in the figure.
Between the gland-cells are long and very attenuated interstitial
cells, of which the outer ends! are expanded and produced to
form a cover over the outer ends of the gland-cells. The nuclei
of the interstitial cells lie in their expanded outer ends. I could
find no trace of cilia. In nearly every specimen examined the
mantle-cavity was full of a sticky gelatinous mass secreted by the
hypobranchial gland.

From what precedes it is evident that the Helicinide are a very
homogeneous group, so far as their anatomical characters are
concerned, and that such differences as I have been able to detect
are of very little assistance in classification, the resemblances and
differences in one set of organs suggesting one class of affinities,
those of another set of organs suggesting another class. Thus,
taking the arrangement of the coils of the intestine as a cr iterion,
we should place Alcadia, Or obophana, and Paleohelicina alse
together, with Zucidella related but somewhat apart : Hutrochatella
would go with Aphanoconia to form a distinct group. But if we
took the characters of the female gonaducts as a criterion we
should get a different arrangement. WHutrochatella would stand
nearest to Alcadia, with Aphanoconia more distantly related :
Paleohelicina with Orobophana would form a distinct group.

The nervous system is so similar in all the species that it affords
a very slight guide, but, as far as the cerebral ganglia give any
clue, the genera would fall into an arrangement consistent with
their geographical distribution; Alcadia, Lucidella, and Hutro-
chatella forming one group; Paleohelicina, Orobophana, and
Aphanoconia another. Finally, the radular characters, which
have attracted a considerable share of the attention of systematists
and in my experience afford the most reliable and readily

MORPHOLOGY OF THE HELICINIDA. 797

recognizable marks of distinction between different species,
give a totally different result. Taking these as a criterion,
Eutrochatella stands well apart: the remaining genera show a
sufficient amount of similarity to justify our placing them in a
single group, in which Paleohelicina stands nearest to Alcadia ;
Aphanoconia is closely related to Palewohelicina; Orobophana, 3
while showing relationship to the two last named, has distinctive
characters which keep it somewhat apart; and Lucidella, while
showing relationship to <Alcadia, has undergone modifications
which, in one feature at any rate, resemble those which distin-
euish Orobophana from Palwohelicina.

On the whole, the radular characters afford the safest clue to
affinity, agreeing as they do with the conclusions founded on
conchological characters and on geographical distribution.

In all the Helicinide, so far as is known, the radular formula
may be represented as co .1(3+1+3).1.0. The Proserpinide
have a somewhat different, and the Hydrocenide a very different,
formula, but they need not be considered here.

As different authors use different names in deseribing the teeth
of Neritoid gastropods, I must define my terms before proceeding
further. Of the teeth included in brackets in the formula, I call
the single tooth in the centre the median; the three teeth on
either side of it the admedians. he large tooth on either side of
the admedians | call the lateral, and the numerous teeth to the
outside of these the marginals. In Hutrochatella, as Troschel has
shown, the lateral teeth are relatively very large and of charac-
teristic shape, being mushroom-shaped, with the top of the pileus
hollowed out to form an articular cavity, into which fits the stalk
of the lateral tooth of the row next in front of it. The edge of the
pileus is entire. I have attempted in fig. 57 (PI. IL): to give
some idea of the very complicated str neues of this tooth in Hutro-
chatella pulchella. It should be noted that it has an external

rocess, or ‘‘ Basalanhang,” which Troschel described as charac-
teristic of the genus Helicina. For the rest, this mushroom-shaped
tooth more nearly resembles in shape the corresponding radular
tooth in the Neritide than is the case in any other Helicinid.
The marginal teeth of Hutrochatella ave simply pointed curved
bars, without denticulations at their free extremities. In the
genus Helicina Lamarck (non sensu restricto Wagner) the laterals
are not pileiform, but consist of a stout median portion which I
shall call the “‘ stalk”; from the inner side of this a more or less
broad aliform plate projects obliquely forward; the anterior
border of this plate is thickened, recurved, and bears a number of
denticulations, varying from 7 to 12 in number in the different
species that I have studied. Attached to the outside of the stalk
by an imperfect joint is the pointed external process (the
‘“‘ Basalanhang ” of Troschel), and the top of the stalk is excavated
to form an articular cavity for the hinder end of the stalk of
the corresponding tooth in the row next preceding. Also, in all
species of Helicina the marginal teeth have broadened recurved

Proc, Zoot, Soc.—1911, No. LY. 5o

798 PROF. G. 0. BOURNE ON THE

anterior extremities bearing denticulations. Thus, there is a
considerable difference between Hutrochatella and Helicina, but the
gap is bridged over by 7’rochatella chrysochasma, 1a which, accord-
ing to Troschel, the lateral tooth is pileiform, but with an oblique
anterior border bearing from 7 to 9 denticulations, and whereas
the proximal marginal teeth are simply pointed as in Hutrochatella,
the more distal marginals bear denticulations, increasing from one
to four innumber. I haveshown that anatomically Hutrochatella
bears the closest resemblance to Alcadia, differing only in the
arrangement of the coils of the intestine.

The genus Alcadia is characterized by a notch separating the
peristome of the shell from the columella. It is stated (Fischer,
‘Manuel de Conchyliologie, p. 795) that the operculum has a
dentiform process: I can only say that I cannot find a trace of
any such structure in A. palliata and A. hollandi. In all other
respects Alcadia is similar to the point of identity to Helicina.
Troschel declares that the only recognizable difference between the
radule of Aleadia and Helicina is in the form of the median tooth,
a character of very little value, for, as I shall show, the shape of
this tooth varies from species to species. But Troschel’s figure of
the radula of Alcadia is not very exact. J have given in Pl. XL.
fig. 58, a to f, large scale drawings of the teeth of A. hollandi;
those of 4. palliata differ only in minute particulars. Comparing
these with the drawings of the radule of Lucidella, Palecohelicina,
Orobophana, and Aphanoconia (Pls. XL.-XLII. figs. 59-65), it
will be seen that the first admedian tooth of Alcadia has a
characteristic shape, being subquadrangular in outline, with its
anterior outer angle produced into a knob on which are borne
four blunt denticulations. The large lateral has a short stalk,
hardly projecting behind the origin of the aliform plate: the
latter is large, expanded, bearing seven stout but blunt teeth on
its recurved anterior margin. The articular excavation is very
shallow: the external process long and pointed.

Of the other radule figured, those of Palwohelicina (Pl. XLI.
fig. 60) and Aphanoconia (Pls. XLI. & XLII. figs. 62-65) bear the
closest resemblance to Alcadia. Both these genera were included
in Helicina Lamarck, and have only recently been separated by
Wagner. In them the lateral teeth have the same general shape
as in Alcadia, but the stalk is longer, the aliform plate less
expanded, the denticulations on its anterior border vary in size,
shape, and number. The first admedian tooth is very similar in
the two genera, and differs from that of Aleadia. The median
tooth is very variable in size and shape. The similarity of the
radulee of Palcohelicina and Aphanoconia is remarkable, and leads
one to doubt whether Wagner is justified in placing these forms
in different genera. On the other hand, Orobophana (Pl. XLI.
fig. 61) is distinct: its lateral. tooth is that of Paleohelicina, but
the first admedian is relatively large, acutely triangular, its
anterior border thickened but without dentieulations; it is
feebly corneous and nearly transparent, suggesting that it is in

MORPHOLOGY OF THE HELICINID®. 799

course of disappearance. The median tooth is heart-shaped,
minute, and similarly feebly corneous. Lweidella (Pl. XL. fig. 59)
is quite distinct in radular characters: in the lateral teeth the
stalk is practically obsolete, the bulk of the tooth consisting
mostly of the aliform process with its thickened denticulate
border, which is continued posteriorly into the articular knob.
There is no anterior articular excavation, but a thin triangular
external piece which serves to support the articular knob of the
tooth of the row next in front, and for the attachment of the
external process. The third admedians are of the usual petaloid
shape; the second admedians stout, triangular, with a thickened
curved anterior edge, bearing on its outer surface a small minutely
denticulate trenchant process. The first admedians are rather
large, but feebly corneous, with a thickened anterior non-denti-
culate border; they have been modified in a manner analogous
to what has been observed in Orobophana. The medians are
broadly heart-shaped, feebly corneous, nearly divided into two by
a deep median anterior notch.

Summing up these details and taking into comparison Troschel’s
figures, which are mostly of species of. Helicina sensu restricto of
WwW agner, and confining our attention to the lateral tooth, which is
the largest and obviously of most functional importance in the
Ner itacea as well as in the Helinacea, we see that there is an easy
transition from LHtrochatella to Alleadia that the lateral tooth
of Alcadia is of the form characteristic of the Helicinide in
general, but shows a tendency to a reduction of the stalk, which,
as Troschel has shown, is common to many American and West-
Indian species. This tendency is exhibited in an extreme form
by Lucidella. But in the Pacific and Oriental genera the stalk
and the articular excavation connected with it are well developed.
But the lateral tooth of Hutrochatella bears an extremely close
resemblance to that of the Neritide, and there is this further
resemblance, that the first admedian tooth, which is of very large
size in the Neritide, is relatively of much lar ger size as compar ed
with .the second and third admedians in Hutrochatella than in
any other Helicinid. If such characters can be relied upon as a
guide to affinity, Hutrochatella is the most closely related among
the Helicinidee to the Neritoid ancestor of the group. From
Eutrochatella forms have been derived: on the one hand, the
Proserpinidee, which also have a large pileiform lateral fond: :
on the other hand, Helicina. We may infer that the earliest
Helicine retained the stalk and articular excavation which are
such marked features in the pileiform lateral tooth of Hutro-
chatella and Proserpina.

The forms which, as suggested in the earlier part of this paper,
were transported bys some unknown means across the Pacific
Ocean to the Philippines must have possessed these features
and transmitted them unchanged to their descendants which
now inhabit the Oriental and Indo-Pacific regions. But in
America and the West Indies there has been a tendency, more

55*

800 PROF. G. G. BOURNE ON THE

fully realised in some species than in others, to’a reduction of the
stalk and articular cavity, this reduction being shown to a slight
degree in Alcadia, to a marked degree in Laucidella.

In all the Helicinids there is a tendency to the reduction of
the central and admedian teeth: this tendency is shown in a
marked deeree in Lucidella and Orobophana, but must have
reached its present degree independently in these two genera.
The reduction is carried to an extreme degree in Hydrocena, in
which the second and third admedians have disappeared; the
median and first admedian are present, but in a rudimentary
condition, and the laterals are reduced to mere rods of no great
size. But I am disposed to think that the “Hydrocenide must
have branched off from the Neritoid stock independently of the
Helicinidee. They retain many primitive features, as Thiele has
shown, among others the process of the operculum which is quite
Neritoid in character, and their geographical distribution favours
this view. Mydrocena is contined to the marine littoral of
Dalmatia; Georissa lives at considerable altitudes on the Khasi
Hills in India. It is by no means improbable that pulmonate
forms may have been developed more than once from such
animals as the’Neritide, which show a predilection for migrating
as far as possible out of the water, and for the rest of it, the most
that can be said in favour of uniting the Hydrocenide with the
Helicinidee is that both display strong Neritoid affinities.

The main result of my researches is to show that in such a
limited group as the Helicinide the systematists are justified in
their methods. The visceral anatomy of all the forms that I
have examined is strikingly similar, and where deviations occur
they are contradictory and of uncertain value. The Helicinide
appear to have inherited an organization with marked Neritoid
characteristics, and to have maintained it, with little or no
change. Presumably that organization is well adapted to the
somewhat narrow range of the conditions of their existence, and
any deviation from it has been checked by the action of natural
selection. But there are a thousand deviations, 1m all directions,
among characters which cannot by any stretch of the imagination
be claimed to be of any importance in the struggle for existence.
Such characters are the texture and coloration of the shell; the
shape of the aperture; the extent and distinctness of the basal
eallus ; the presence or absence of folds at the aperture of the
shell; the presence or absence of a minute notch, such as occurs
in Aleadia; the arrangement of the growth-lines on the oper-
culum. It might be said that the operculum is an important
protective organ and therefore eminently susceptible to the action
of natural selection. But its function is simply to close the
aperture of the shell, and this it does equally efficiently in all
the species that I have examined, the number of these being
much larger than the few available for anatomical study. As
long as the operculum performs this function efficiently mmute
characters, such as the greater or less distance of its nucleus from

MORPHOLOGY OF THE HELICINIDA. 801

the anterior border, cannot possibly determine the question of
the death or survival of the animal. The same reasoning applies
to the variations of the radular teeth : the function of the radula
is to rasp, and any of the modifications shown in figs. 60 to 65
is equally efficient as a rasp. Nobody, I think, would venture
to assert that the minute differences in the four species of
Aphanoconia (figs. 62 to 65) could have had any value in the
differentiation of these species by natural selection.

Asa result of my somewhat elaborately minute studies, I am
driven, and, I confess, somewhat unwillingly driven, to the con-
clusion arrived at by a number of naturalists, that natural
selection is efficient in preserving characters of physiological
importance, but ineffective in producing new species by adding
together numerous minute successive variations. The only
conclusion justified by the facts seems to me to be that the
characters on which systematists rightly rely are of the nature
of deviations or mutations, of no consequence to the well-being
of the animals in which they appear, but inheritable, and there-
fore perpetuated under favourable circumstances by segregate
breeding. The Helicinide, inhabiting narrow areas, and often
segregated in remote islands, afford particularly favourable oppor-
tunities for segregate br eedin, g,

As to how far hase small deviations of functionally unimportant
structures may be due to the influence of external conditions I
do not venture to offer an opinion, but the following fact is
suggestive. Among the shells in the tube containing several
specimens of Viphaiacanae merguiensis was a specimen which
in size, shape, coloration, and marking so exactly resembled the
others ‘that I took it fon a Helicinid (as the colleetor must also
have taken it) and decalcitied it with a view to anatomical
investigation. It proved to be a Helicid, of what genus and
species I cannot say, as I had destroyed the shell and could not

find another specimen.

Among the collection of Helicinide made in the Andaman
Islands and presented to the British Museum of Natural History
by Mr. G. Rogers was a tube containing half a dozen specimens
which differ recognizably in the characters of the shell and
operculum from Aphanoconia andamanica Benson, but are clearly
closely related to that species. I have not been able to refer
them to any named species, and as the radular characters show
it to be distinct from anrdamanica, | describe it as a new species,

as follows :—

APHANOCONIA ROGERSII, sp.n. (PI. XLII. figs. 65-69.)

Shell oblately spheroidal, the surface marked with closely set
radial growth-lines ; colour ight orange-yellow marked with more
or less distinet reddish-brown radial bands ; spire of 47 whorls,
increasing regularly and somewhat rapidly in size, the last whorl
obtusely keeled, the keel produced into a prominent angular

802 PROF. G. C. BOURNE ON THE

projection at the peristome. Aperture semilunar, very oblique,
the outer margin thickened and expanded. Basal callus not
very thick, hardly differing in colour from the rest of the shell,
of rather small extent, its limits clearly defined above, as well as
below. ‘The whole shell deeper in proportion to its breadth than
in A. andamanica and the spire more prominent.

Operculum yellowish white in colour, the calcareous plate
rather thin, the sigmoid curve pronounced, the upper angle
produced.

Radula with small diamond-shaped median tooth; the first
admedian tooth more than twice as long as broad, the anterior
edge incurved and bearing four denticulations ; second and third
admedian teeth each with four denticulations; the lateral teeth
normal, the stalk rather long, the articular excavation deep, the
aliform process bearing seven round denticulations.

Closely as the shell of this species resembles that of A. anda-
manica, a glance at the drawings of the radule of the two species
(figs. 63 & 65) shows that they are distinct.

I must express my obligations to Miss Margaret Poole, both
for helping me in the determination of the different species of
Aphanoconia and for making the drawings of shells and radule

for figs. 62 to 68.

As I have discussed and offered an explanation of the geo-
graphical distribution of the Helinicide without either adopting
or criticizing the theories advanced by Dr. Simroth (7 and 8) on
this subject, I must, in conclusion, make some reply to the friendly
eriticisms that he has published on my paper on the Neritide.
I do not propose, in this place, to discuss the physiological
interpretation that he has given of the different arrangements of
the female ducts in the Neritide. For one thing, I have
obtained some new material and hope soon to publish further
observations throwing fresh light upon the problems to which
he refers: for another, J am inclined to accept much of what he
writes on that part of the subject.

_ But with regard to the ancestry of the Neritide, and with them
the Helicinide, which Dr. Simroth would derive from a pulmonate
stock, and with regard to the homologies that he wishes to establish
between the generative ducts of Neritide and Pulmonata, I am
unable to accept any of his conclusions. To do so would be
to throw the whole fabric of morphological reasoning to the
ground. Dr. Simroth’s views on homologies are largely influenced
by a theory of secular changes in the sea-level produced by a
swinging or “ pendulating” movement of the earth about an axis
which corresponds with the longest diameter of the earth and has
its poles in Sumatra and Ecuador. It is not my present intention
to discuss the difficult astronomical and geological problems in-
volved in the “ Pendulation theory,” and, indeed, I am sure that I
am incompetent to discuss them. The theory may be well founded
or 1t may not: I do not offer an opinion; but be it right or

MORPHOLOGY OF THE HELICINID. 803

wrong, I fail to see that the conclusions deduced from it by Dr.
Simroth are necessary. I will explain as briefly as possible why.
Unless I misunderstand him grievously, and if I do I beg his
pardon, one of Dr. Simroth’s chief conclusions is that, contrary to
the generally accepted doctrine, marine and freshwater animals in
general, the marine prosobranch Gastropods in particular, are
evolved from terrestrial forms which have been forced by the
above-mentioned secular inundations to adapt themselves to new
conditions of life and make their habitat inanother medium. As
the pendulation theory applies to all geological time, if the pre-
cursors of marine Gastropods were terrestrial in habit, we should
find evidence of this in geological deposits. The earliest-known
Gastropods, from the Cambrian to the Devonian, would bear
evidence of their terrestrial life, those found in later deposits
would indicate, in some periods at least, the change from a ter-
restrial to a marine existence. But, in point of fact, the geological
evidence points decisively the other way. In Cambrian, Ordovician,
Silurian, and Devonian deposits we get Gastropods belonging
almost exclusively to the Streptoneurous Aspidobranchia and
Pectimibranchia. There are, it is true, the pteropod-like shells of
the Conularida which, if they are really remains of Pteropods,
would demonstrate the great antiquity of highly specialized forms
of Kuthyneura. But the true systematic position of the Cornu-
larida is at the best doubtful, and it has been urged with much
reason and on high authority that the resemblance between the
shells of these archaic forms and the more modern Pteropoda is
due to parallelism. As so much doubt prevails as to their affinities,
the Cornularida cannot be brought into the argument. The
Aspidobranchiate and Pectinibranchiate Gastropods from these
earlier Paleeozoic deposits are without doubt marine forms. They
subsisted, without any important changes, through the four above-
mentioned geological epochs, and one genus, Plewrotomaria, has
survived to the present day. We know the habits and the
anatomy of Plewrotomaria, and they support in a most remarkable
manner the conclusions derived from an extensive knowledge of
gastropod morphology. On the other hand, with the exception of
Hercynella from the Devonian, undoubted Euthyneura first make
their appearance in the Car ‘bonifer ous. They belong to the
Acteonide and Pulmonata Stylommatophora. The first-named
family is marine, and anatomically displays so many strepto-
neurous characters that it might almost be included in the
Aspidobranchia. Of the Sty lommatophora we get forms hke
Dendropupa and Pyranidula, unquestionably terrestrial species,
and, according to views generally accepted, highly modified and
therefore indicative of a line of lost ancestry probably allied to the
contemporary Actzonide. But these pulmonate forms are few
and of rare occurrence in the Carboniferous, a period in which
the conditions for the preservation of terrestrial and freshwater
forms were particularly favourable. Had numerous Pulmonates
existed at that time their remains must have been more abundantly

804. PROF. G. G. BOURNE ON THE

preserved, Terrestrial pulmonates are still scanty in the Permian
and Trias, and only begin to show a considerable increase in the
Jurassic and Cretaceous. I need not labour the point further.
Clearly, paleontological evidence does not favour Dr. Simroth’s
theory of the orgin of marine from terrestrial Gastropoda.

But let us suppose that paleontological evidence may be ignored
on account of the imperfection of the geological record, and that
the Pendulation theory is so well supported by other evidence as
to compel us to give credence to Dr. Simroth’s doctrines as to the
origin of marine from terrestrial Gastropods. The Helicinide
are terrestrial and pulmonate. I have shown, and in so doing
have only corroborated the opinion of all other observers, that
they are Neritoid in almost every feature of their anatomy. If
the marine and fluviatile Neritids were to be derived from a ter-
restrial and pulmonate form, one would suppose that that form
must have been Helicinid in character, for the affinities between
the two groups are so very obvious. But Dr. Simroth does not
discuss this possibility. Making reference to Ostracolethe,
Hyalimax, Limax, and Arion, all highly specialized recent
Pulmonates, he boldly derives the Neritide from the Stylomma-
tophora, relying largely upon the supposed homology of their
generative ducts. This homology I do not admit: a resemblance
there i is, but not a close one, and, even if it were closer than it
actually i is, L should place very little reliance on the anatomy of
the generative ducts as indicative of relationship between groups
differing widely in all other respects. In the different phyla of
invertebrated animals the generative ducts are notoriously variable
in character. In the Platyhelmia, for example, their variety is
bewildering. Within the phylum Mollusca there are many in-
stances of variability and also of deviations which must have been
independently acquired but are in the same direction, as, for
instance, in the Doridomorpha and Elysiomorpha. The re-
semblances, such as they are, between the generative ducts of the
monecious Pulmonata and the dicecious Neritidee are just what
one might expect to find in animals in which a common plan of
organization, to wit a gastropod organization, is modified im
accordance oath etnies physiological requirements. The differ-
ences are of amply sufficient magnitude to betray a difference of
origin. In other words, the complex g gonaducts of Neritide and
Pulmonata are independently acquired structures, and such
resemblances as they display are due to parallelism.

T have already referred to the anatomy of the Pleurotomariide,
a family which existed in the Cambrian and survives to the present
day. Thanks to Bouvier and M. F. Woodward, we are well
acquainted with the anatomy of Plewrotomaria, which affords a
striking confirmation of the reliability of sound morphological
reasoning. Before Pleuwrotomaria had been studied, comparative
anatomists, as the result of extensive investigations of gastropod
structure, had come to an agreement concerning numerous marks
of primitive organization in the group. When this survivor from

MORPHOLOGY OF THE HELICINIDA. 805

the Paleozoic age came to be examined, all these marks were
found, some of them in a more pronounced degree than in any
other known Gastropod, and in no system of organs were these
marks more conspicuous than in the nervous system, the im-
portance of which Dr. Simroth seeks to minimize. Among these
marks may be enumerated—a cerebral commissure situated far
forward on the pharyngeal bulb; a distinct labial commissure ;
elongated and scalariform pedal nerve-centres ; a long crossed
visceral commissure; two auricles to the heart; the ventricle
lapped round the rectum; a rhipidoglossate dentition. Other
characters might be enumerated, but these suffice for the present
purpose. All these characters are absent in the Pulmonata : all
of them are present in the Nevritide. Moreover, by discovering
the oviduco-celomic funnel, I was able to demonstrate, beyond all
reasonable doubt, the homology of a part of the gonaducts to the
right kidney of Pleawsotomaria and other rhipidoglossate Aspido-
branchs, a homology which Thiele had already asserted on other
grounds. Now it is quite clear that, if structural resemblance is
of any value as a guide to affinity, we have a choice between two
alternatives. Bither the Neritide, to which we must add the
Helicinide, are descended from Aspidobranch ancestors, which
they resemble in all the points enumerated above, and have in-
dependently acquired genital ducts superficially similar to those
of Pulmonata; or,as Dr. Simroth will have it, they have descended
from stylommatophorous Pulmonata, have preserved the characters
of the genital ducts of the latter group, but have independently
acquired all the other characters enumerated above, characters
possessed by no Pulmonate, but invariably present in those
Aspidobranchs from which, on Dr. Simroth’s showing, the
Neritide are not descended. I am not quite sure whether he
would go so far as to assert that the remaining Aspidobranchs
possess those characters because they are descended from the
Neritide. To make such an assertion would, indeed, be flying in
the face of all reasoned opinion on this subject, and would amount
to a declaration that the geologically more recent Pulmonates are
the parents of their predecessors of Cambrian age !

T submit the alternative to the judgment of my readers, and in
doing so beg leave to enter a protest against the srowing tendency
to throw over long-established and carefully reasoned conclusions
founded upon morphological evidence, because of thei uncon-
formity with some new and as yet insufficiently tested hypothesis,
or because they do not help in the solution of certain limited
problems. I was quite aware, when I discussed the subject, that
the geographical distribution of the Neritide was a puzzle, and
that L had failed to find a solution to it. The distribution of the
Helicinide is scarcely less puzzling and awaits a final solution.
But with all respect for Dr. Simroth’s authority and deserved
reputation as a zoologist, | submit that the solution that he offers
is improbable, raises a crop of other puzzles, and throws
morphology into confusion.

806

(1)

(2)

(3)

(4)
(5)
(6)
(7)
(8)
(9)

(10)

(11)
(12)

PROF. G. C. BOURNE ON THE

List or LireERATURE REFERRED TO.

Amauprur, A.—La partie antérieure du tube digestif et la
torsion chez les Mollusques gastéropodes. Ann. des Sci.
Nat. (8) vii. 1898, p. 1.

Bourne, G. C.—‘‘ Contributions to the Morphology of the
Group Neritacea of Aspidobranch Gastropods.—Part I.
The Neritide.” Proc. Zool. Soe. Lond., 1908.

Bouvier, EK. L.—‘‘ Systeme nerveux, Morphologie générale
et Classification des Gastéropodes Prosobranches.” Ann.
des Sci. Nat. (7) i1., 1887.

IsENKRAHE, C.—‘‘ Anatomie von Helicina titanica.” Arch.
f. Naturgeschichte, xxxiil., 1867, p. 50.

JHERING, H. von.—Vergleichende Anatomie des Nerven-
systems und Phylogenie der Mollusken. Leipzig, 1877.
Kopett, W.—Studien zur Zoogeographie. Wiesbaden, 1897.
Srurora, H.—‘“ Neuere Arbeiten iiber die Morphologie und
Biologie der Gastropoden.” Zool. Zentralblatt, Bd. xvi.,

ISOS),

SimrotH, H.—“The Anatomy of the Neritide.” Proc.
Malacol. Soc. Lond., ix. 1910, p. 27.

THIELE, J.—‘‘ Die systematische Stellung der Solenogastren
und die Phylogenie der Mollusken.” Zeitschr. f. wiss.
Zool., xxii. 1902, p. 249.

Ture, J.—“ Uber die Anatomie von Hydrocena cattaroensis
Pf. Abh. d. Senckenbergischen Naturf. Gesell., xxxi1.,
1910.

TroscHEL.— Das Gebiss der Schnecken. Berlin, 1856-1863,
Bd. 1. pp. 75-85.

Waener, A. J.—‘“ Helicinenstudien.” Denkschr. d. kais.
Akad. d. Wissenschaften, Wien, Bd. Ixxvii., 1905, and
Ixxviu., 1906.

(A complete list of the literature of the Neritoidea is given in
my previous paper [2 ]).

EXPLANATION OF THE PLATES.

Pruates XXX.-XLII.
Lettering in all the figures.

an. Anus. con. ped. Cerebro-pedal connective.
ao. Aorta. | con. pl. Cerebro-pleural connective.
ap. mg. Aperture of hypobranchial ce. ph. Lateral pharyngeal cartilage.
gland. div. Diverticulum of male gona-
Au. Auricle. duct.
bue. Buccal cavity. E. Hye.
c@. Cxcum of ootype. F. Foot.
cl. Cloaca. g. buec. Buccal ganglion.
e.m.l. Left columellar muscle. g. cer. Cerebral ganglion.
ce.m.r. Right columellar muscle. Gd. Gonaduct.
com. bucec. Buccal commissure. gl. pd. Pedal gland.
com. cer. Cerebral commissure. gi.r. Glandular ridge on floor of
com. lab. Labial commissure. the csophagus.
com. pd. Pedal commissure. g- pl. Pleural ganglion.

com. pl. Pleural commissure. Int. Intestine.

MORPHOLOGY OF THE HELICINIDA, 807

Ti. Liver. oot. Ootype.
li. d. Liver-ducts. Op. Opercultum.
M. Mantle. ot. Otocyst.
M. c. Mantile-cavity. ov. Ovary.
m. g. Hypobranchial gland. Pe. Pericardium.
n.cm.l. Left columellar nerve. ped. Pedal nerve-cords,
n.cm.r. Right columellar nerve. ph. Pharynx.
n. gen. Genital nerve. R. Rectum.
n. gl. p. Nerve to pedal gland. rd. Radula.
n. lpb. Labio-proboscidean nerves. rd. s. Radular sac.
m. oc. Ocular nerve. 7. p.c. Reno-pericardial canal.
n. op. l. Left opercular nerve. r. s. Receptaculum seminis.
n. op. r. Right opercular nerve. s. gl. Salivary gland.
n. ot. Otocyst nerve. Sn. Snout.
n. pal.-c.l. Left pallio-columellar nerve. sp. d. Sperm-duct.
n. pal.-c.r. Right pallio-columellar nerve. spz. Spermatozoa.
n. pal.l. Lett pallial nerve. St. Gisophageal moiety of
n. pal.r. Right pallial nerve. stomach.
n. par.l. Left parietal nerve. S¢!. Pyloric moiety of stomach.
n. par.r. Right parietal nerve. é. Tentacle.
n. sb.i. Subintestinal nerve. | é. s. Terminal sac of male gona-
n. ten. Tentacular nerve. duct.
od. Oviduct. Ur. Ureter.
od', Descending limb of V-shaped Ur.p. Uropore.
portion of oviduct. V. Ventricle of heart.
od*. Ascending limb of V-shaped vag. Vagina.
portion of oviduct. vag- ap. Opening of vagina into
od. c. a. Anterior odontophoral carti- mantle-cavity.
lage. | vg. s. Vaginal sae.
od. c. p. Posterior odontophoral carti- | V.G1. First visceral ganglion.
lage. | V.G*. Second visceral ganglion.
oe. (sophagus. | v. pal. Pallial vein.
ce. p. (Esophageal pouch. v. pst. Posterior pallial vein.
0.@.p. Opening of esophageal pouch | v. ren. Afferent renal vein.
into cesophagus.

Fig. 1. Left side view of Alcadia palliata. The mantle has been cut through close
to the left columellar muscle and turned back to expose the interior of the
mantle-cavity, the pericardium has also been opened.

Fig. 2. Dorsal view of the buccal cavity, pharynx, and anterior part of the
esophagus of Alcadia palliata: the buccal cavity, pharynx, and part of
the cesophagus have been laid open; gl.r., glandular ridge on the floor
of the cesophagus.

Fig. 3. A dissection: of the cesophagus and pharynx of Aleadia palliata, seen from
the left side.

Fig. 4. A horizontal section through the pharynx and buccal cavity of Alcadia
hollandi. ,

Fig. 5. The odontophoral cartilages of Aleadia palliata, viewed from below.

Fig. 6. The odontophoral cartilages of Hutrochatella pulchella, viewed from above.

lee Yo AN seo of the stomach of Alcadia palliata, viewed from the ventral
side.

Fig. 8. A portion of the epithelium of the stomach of Alcadia hollandi, showing
glandular and ciliated cells. about 960.

Fig. 9. Part of a section through the prominent ridge in the stomach of Alcadia
hollandi, < about 600; cu., the thick cuticle covering the ridge.

Fig. 10. The alimentary tract of Alcadia palliata, showing the arrangement of the
intestinal coils.

Fig. 11. Alimentary tract of Alcadia hollandi.

Fig. 12. Alimentary tract of Pale@ohelicina ide.

Fig. 13. Alimentary tract of Orobophana ponsonbyi.

Fig. 14. Alimentary tract of Zwcidella awreola.

Fig. 15. Alimentary tract of Hutrochatella pulchella.

Fig. 16. A horizontal section through the upper part of the visceral mass of
Alcadia hollandi. Figs. 17 to 22 are drawn from the same series of
sections.

Fig. 17. A section somewhat lower down, showing the origin of the ureter, Ur.,

from the kidney.

808 ON THE MORPHOLOGY OF THE HELICINIDA,

Fig. 18. A section still more ventral than the above, showing the deepest part of the
pericardium and its relation to the kidney.

Fig. 19. A more ventral section passing through the reno-pericardial canal,
Vp.c.

Fig. 20. A ee section passing through the ventral part of the visceral mass.

Fig. 21. Part of a section slightly ventral to that drawn in fig. 19, showing the
uropore, Uy.p., opening into the mantle-cavity.

Fig. 22. A section through the reno-pericardial canal. Magnified about 600.

Fig. 23. Part of a section passing through the uropore of Lucidella awreola and
showing the character of the renal epithelium. X 960.

Fig. 24. A diagram reconstructed from the series of sections drawn in figs. 16 to 20,
showing the relations of the kidney, ureter, stomach, pericardium, and
mantle-cavity.

Fig, 25. The genital ducts of Aleadia hollandi 2. 'The lower half of the figure is
drawn as it appears when viewed by transmitted light, the upper part as
seen by reflected light. In this and in figs. 26-29, 40, and 41, the
gonaducts are represented as seen from the ventral side, after the wall of
the mantle-cavity has been cut through by a dorsal incision and the
rectum and gonaducts turned over to the right side of the animal.

Fig. 26. The genital ducts of Hutrochatella pulchella 9 .

Fig. 27. The genital ducts of Aphanoconia merguiensis 2.

Fig. 28. The genital ducts of Paleohelicina ide ¢.

Vig. 29. The genital ducts of Orobophana ponsonbyi & .

Fig. 30. A longitudinal section through the genital ducts of Alcadia hollandi ?,
passing through the aperture of the hypobranchial gland.

Fig. 31. A longitudinal section from the same series, showing the aperture of the
vaginal duct.

Fig. 32. Another section from the same series, showing the origin of the oviduct
from the ovarian chamber.

Fig. 33. Another section from the same series showing the connection of the vagina
with the vaginal sac and ootype.

Fig. 34. Another section from the same series showing the opening of the oviduct, od.,
into the descending limb of the V-shaped tube, od.’

Fig. 35. Another section from the same series showing the receptaculum seminis
opening into the ascending limb of the V-shaped tube.

Fig. 36. Glandular epithelium from the wall of the ootype of Alcadia hollandt.

Fig. 37. Ciliated epithelium and spermatozoa from the receptaculum seminis of
Aleadia hollandi.

Fig. 38. A section through the oviduct of Alcadia hollandi. Highly magnified.

Fig. 39. An epithelial ridge from the bilobed cecum of the ootype of Orobophana

onsonbyi.

Fig. 40. ee of the genital ducts of Aleadia hollandi 6.

Fig. 41. A similar view of the genital ducts of Aphanoconia gouldiana 6.

Fig. 42. A drawing made from a combination of several serial longitudinal sections
through the genital ducts of Hutrochatella pulehella 8, showing the
narrow diverticulum, &.7., which may possibly represent the vagina of the
female and therefore be the homologue of the right kidney-sac.

Fig. 43. A dissection showing the pedal, pleural, and visceral nerve-centres in
Aleadia palliata, with the principal nerves issuing from them. The
cerebral ganglia have been removed. The dissection is made from the
dorsal surface; the foot, as is usual in contracted specimens, is turned
forward and lies in front of the head with the sole uppermost; the walls
of the head and the mantle have been cut away, and the visceral mass has
been dissected as far as is necessary to show the course of the subintestinal
nerve and its branches.

Fig. 44. The nerve-centres and principal nerve-trunks of Alcadia hollandi, viewed
from the right and above. The drawing was made with the camera
lucida, after removal of the nerve-centres from the body.

Figs. 45-52. A series of transverse sections through the pleuro-pedal nerve-centres of
Alcadia hollandi, showing the principal tracts of nerve-fibres in the fused
pedal, pleural, and subintestinal ganglia. For a full description of these
figures, see the text, p. 789. The position of the pedal gland is indicated
in fig. 45.

Fig. 68. The left cerebral ganglion of Paleochelicina ide, viewed from the inner
surface. The ganglion is stained with Mayer’s hemalum and drawn by
transmitted light ; 1, 2, 3, 4, the four labio-proboscidean nerves.

Fig. 54. A similar preparation of the left cerebral ganglion of Alcadia palliata.

ON THE PALATABILITY OF SOME BRITISH INSECTS. 809

Fig. 55. A drawing of a section showing the position, structure, and nerve-supply of
the opercular organ of Hutrochatella pulchella.

Fig. 56. A portion of the epithelium of the hypobranchial gland of Alcadia
hollandi. Wighly magnified.

Fig. 57. A left lateral tooth from the radula of Hutrochatella pulchella. Highly
magnified.

Fig. 68. Radular teeth of Aleadia hollandi, highly magnified: a, median; 6, c, d,
first, second, and third admedians of the left side; f; one of the marginals
or uncini; é, a lateral tooth of the right side showing the stalk, stk., the
aliform internal plate, al.p., the articular excavation, art., and the
process, ext.p.

Fig. 59. Three rows of teeth from the radula of Lucidella aureola. In this and
the following figures only the proximal members of the marginals are
indicated.

Fig. 60. Two rows of teeth from the radula of Palg@ohelicina ide.

Fig. 61. Two rows of teeth from the radula of Orobophana pachystoma ponsonbyi.

Fig. 62. Two rows of teeth from the radala of Aphanoconia gouldiana.

Fig. 63. Two rows of teeth from the radula of Aphanoconia andamanica.

Fig. 64. Two rows of teeth from the radula of Aphanoconia merguiensis.

Fig. 65. Two rows of teeth from the radula of Aphanoconia rogersii.

Fig. 66. Shell of Aphanoconia rogersii.

Fig. 67. Shell of the same species, showing the aperture.

Fig. 68. Shell of the same species, viewed from above.

Fig. 69. Operculum of Aphanoconia rogersii, viewed from the inner or ventral side.

36. On the Palatability of some British Insects, with Notes
on the Significance of Mimetic Resemblances. By
R. I. Pocock, F.R.S., F.L.S., F.Z.S., Superintendent
of the Society’s Gardens and Curator of Mammals.
With Notes upon the Experiments. By Pict. E. B.
Poutton, F.R.S., F.Z.S.

[Received and Read May 9, 1911.]}

INTRODUCTION.

At the request of Prof. E. B. Poulton, F.R.S., I undertook, in
the summer of 1909 and again in that of 1910*, to make a series
of experiments in the Zoological Gardens to test the palatability
of various species of British Insects. Much of the material was
sent to me by Dr. G. B. Longstaff from Morthoe in Devonshire.
Some I received from Prof. Poulton himself or from friends of
his. A few species I added on my own account; notably the
stick insects and the ants, of which we had an abundant supply
in the Insect House in the Gardens. Those that I supplied
I identified myself. The rest were in all cases named by the
senders. To the insects Dr. Longstaff added a number of slugs,
which were identified, I understand, by Mrs. Longstaff.

Since the majority of the experiments were made with English
Insects, it is regrettable that English, or at all events Palearctic
birds, were, for the most part, unavailable for the tests. There
were two reasons for this. In the first place, Palearctic insecti-
vorous birds were not strongly represented in the Society’s

* Records of a few experiments made in 1911 haye been incorporated in the text,

810 MR. R. I. POCOCK ON THE

collection. Im the second place, those that were in the Zoological
Gardens at the time were, in most cases, kept in a very large
flight aviary with plenty of cover in the way of shrubs, repre-
senting their natural environment as nearly as possible. Never
having been tamed by confinement in small cages, they were too
shy to come to the bars to take insects from my hand and too
seared to notice them if I entered the aviary. Once or twice
I tried the experiment of liberating butterflies in this aviary ;
but the frequency with which they escaped through the wire
mesh and were wasted for the purpose in hand, induced me to
abandon further experiments of that kind.

This reference to the shyness of birds in captivity brings me
to another of the limitations under which I was working. I
was forced to restrict my attention to particular birds, tame
enough either to take insects directly from me or sufficiently
accustomed to the presence of human beings in the aviary to
capture liberated insects in spite of my close proximity. If I
put the insects through the bars, myself standing outside, they
were either seized one after another by the boldest bird in the
place, or were carried by a timid bird to the back of the com-
partment, where | could not watch what befell them. I was
compelled, therefore, to be inside the bars. Since, moreover,
it was practically impossible to watch more than one bird at a
time, I was precluded from the method of experimenting with
the shyer specimens by giving insects to the bolder ones to
distract and monopolize their attention. Thus it comes about that
the same species appear over and over again in the experiments
below recorded, while many insectivorous birds, that might have
been tried but for their shyness, are omitted.

Two facts struck me very forcibly at an early stage of the
experiments. The first was the exceeding keenness of the birds
for the insects brought to them. This was no doubt due in a
measure to our inability in the Gardens to feed the birds on
living insects other than mealworms. ‘The living prey was
evidently a great treat to them; and over and over again I was
impressed with the persistence shown by birds in persevering
with insects that were obviously not to their liking, returning to
the morsels repeatedly as if food of such a nature was too good to
be wasted. From this I think it may be inferred that in a state
of nature hungry birds will eat nauseous insects which in times
of plenty they will reject after tasting, or will not take the
trouble to catch them if they have previously learnt their distaste-
fulness by experience. Furthermore, it is quite clear that the
plain record of an insect being eaten is no proof of its palatability.
Better evidence on this head is supplied by the behaviour of
the bird towards it. After a little experience in this matter, I
was able to satisfy myself at all events as to the approximate
correctness of my interpretation of the bird’s actions, and to
judge thereby of the comparative palatability of the insects they
tasted.

PALATABILITY OF SOME BRITISH INSECTS 811

The second fact has an important bearing upon the criticism
sometimes advanced against the theory of warning coloration
and mimicry as applied to butterflies, namely, that birds under
natural conditions are seldom seen to eat these insects.* Hence it
has been inferred that birds cannot be reckoned as serious enemies
of buttertlies. Whatever may be the explanation of the cireum-
stance, I am tolerably sure, from the behaviour of the two classes
of animals when pitted against one another, that the inference
drawn therefrom is erroneous. The insectivorous birds in our
aviaries seemed to know at once what the butterflies were; they
were on the alert the moment one was liberated and pursued 1t
with determination and precision, following its every turn and
twist, and either catching it upon the wing or pouncing upon it
after settling. It is true that this predatory deftness.may have
been acquired in relation to the chase of insects other than
Lepidoptera; but unless the birds recognised butterflies in
general—a group which cannot be mistaken for other insects—
as part of their natural prey, it is difficult to understand their
eager excitement at the sight of those I offered them.

Again, unless the species of butterflies used for the experiments
are, or were in the past, habitually preyed upon by birds,
whence comes the extraordinary skill the liberated specimens,
when undamaged or inexhausted by confinement, displayed in
dodging the swoop of the birds in mid-air? Having repeatedly
seen the aim of the pursuing bird baftled by the evasive twist
of the butterfly, I cannot doubt that the insect’s behaviour was
prompted by the instinct to escape an habitual enemy of its
species, of the same class and with the same predatory methods.
It cannot, I imagine, be seriously claimed that escape from the
upleap of insectivorous mammals, lizards, or frogs has been a
factor of sufficient importance in survival to be reckoned with
in this connection; and, a fortiori, the modernness of the
invention of the entomologist’s net puts this instrument of
capture out of court for consideration. The evidence, therefore,
seems to me to afford the strongest support to the conclusion
that the power to dodge in mid-air and the instinct to put it
in force have been fostered to subserve no other purpose than
the evasion of swift-winged insectivorous foes. Perhaps predatory
Pompilide must be regarded as a possible auxiliary influence ;
but apart from these hymenoptera, I can think of no enemies but
birds likely to have persecuted butterflies on the wing to the
extent presumably necessary to have guided their evasive tactics
to the pitch of proficiency they now exhibit.

Whatever be the value of this suggested explanation of the
facts, the facts themselves remain as I have stated them :—
(1) Caged insectivorous birds which, so far as is known, have never
been fed in captivity upon butterflies, are at once excited by

* Twice I have seen sparrows, which are not typically insectivorous, chase white
butterflies in London. Two birds acting in concert were successful on the first
occasion ; one singie-handed failed on the second occasion,

812 MR. R. I. POCOCK ON THE

their appearance, chase them with eager speed, catch them in
mid-air with precision, and eat them or taste them with avidity.
(2) Pursued butterflies when overtaken often avoid the birds,
not once only but twice or three times, by sudden turns up or
down to right or left.

Those who hold, on the negative evidence above stated, that
birds are not to be reckoned as serious enemies of butterflies,
must be called upon to sapply some explanation other than that
above proposed of the marked reactions between these two
classes of animals when brought into contact with one another,
and to show reason why what takes place in the aviary may
not be regarded as indicative of similar occurrences in nature.

With regard to the experiments on mimicry, especially those
made with Volucella bombylans and Bombus hortorwm, 1t appears
to me that they satisfy all that the theory, as propounded by
Bates, demands. They fully confirm Prof. Lloyd Morgan’s experi-
ments on birds, with the drone-fly (Hristalis) and the honey-bee
(Apis mellifica), as well as those with the banded and uncoloured
slips of glass holding respectively meal adulterated with quinine
and meal untampered with.* They show that several species
of birds, after learning by experimental tasting that Lombus
hortorum is unpalatable, refused to touch Volucella bombylans.

Other items of interest that may be briefly alluded to are the
experiment demonstrating, at least in the instance tried, the
attractive nature of the ocelli on the wings of the peacock butterfly
(Vanessa io); the experiments showing that Yormica rufa is not
protected from mammals and birds by its acid taste; that the
black members of the Carabide and Ocypus olens are unpalatable
to the ground-feeding mammals they were offered to; that
Coccinella T-punctata and the Telephorid beetle (? Rhagonyche
fulva)—belonging to families of beetles which are common
objects of mimicry in the tropics—are distasteful to nearly all
mammals and birds.

At the end of the part of the paper describing the experiments
made, I have added, at Dr. Longstaft’s suggestion, for the informa-
tion of those unfamiliar with the habits and distribution of the
mammals, birds, and reptiles to which the insects and other
invertebrates were offered, a list of the species of the former
groups giving a few particulars on those points.

Finally I have to thank Prof. Poulton for kindly annotating
the paper before it went to press, and for explaining more fully
than I could do the bearing of some of the results on the
theories of mimicry and of the connection between palatability and
coloration. I am also indebted to Commander J. J. Walker, R.N.,
for kindly giving me the scientific names of the Lepidoptera.

* Animal Behaviour, pp. 164-165, 1900.

PALATABILITY OF SOME BRITISH INSECTS. 813

THe EXPERIMENTS

MOLLUSCA.
(SLues.)
Large Black Slug (Arion ater).

Sept. 24,1910. Two taken and eagerly eaten bytwo Meerkats,
who wiped them down with their paws and rubbed them in the
sand apparently to remove the slime. -

Oct. 26, 1909. One given to Black-winged Grackle was
eaten.

One (larger specimen) given to the same bird was abandoned ;
offered to Sulphury Tyrant, but the bird would not touch it;
offered to Sun-Bittern, was pecked, but not eaten ; carried to a
perch by Harmonious Shrike-Thrush but was soon dropped ;
pecked and shaken about, and much hammered by Abbot’s Rail,
which managed to break the skin of the slug and getting at the
inside ate a large portion, but would not eat the outside.

One taken by Dial Bird which persevered for a long time,
hammering and wiping it in the sand; he was then driven off by
Black-chinned Laughing Thrush, which held the slug in his foot
and ate little pieces of the inside after breaking the skin, but left
the bulk of it.

Dial Bird tried another, but gave it up.

Common Hangnest took one, but left it after a few pecks.

Two offered to Kagu, a kind of Crane or large Rail, were
swallowed entire with very little delay.

One tried by Black-tailed Water-hen which, however, gave it
up; the same specimen given to Leach’s Laughing Kingfisher was
ultimately swallowed entire after being dropped many times.

Arion hortensis.
(Olive-brown Slug with orange-coloured foot.)

Oct. 26, 1909. One eaten by Yellow crowned Hangnest.

T'wo eaten by Dial Bird.

Two refused by Harmonious Shrike-Thrush.

One twice taken from my hand by Harmonious Shrike-Thrush
and dropped both times; but after taking it the third time the
bird ate it.

LTimax maximus. -

Oct. 26, 1909. One given to Harmonious Shrike-Thrush was
tried, but abandoned after one or two pecks. It was then taken
and eaten by a Black-winged Grackle after a great deal of wiping
of the bill.

Sept. 24, 1910. Two specimens tasted by Green Lizard, and
Black-spotted Lizard, but not eaten. The Lizards apparently
disliked the slime, because they wiped their mouths on the stones
after tasting. Both eaten without delay by Glass Snake.

Proc. Zoou, Soc.—1911, No, LVI. 56

814 MR. R. I. POCOCK ON THE

Limax agrestis.

Sept. 24,1910. One eaten after a good deal of pecking about
in the sand by White-crested Jay-Thrush.

Two eaten by a Shama.

One eaten by Kagu.

Tasted but rejected by Fantailed Flycatcher.

Tasted on two occasions by Hoopoe but rejected.

Tasted by Red-vented Bulbul but rejected.

Tasted but rejected by Yellow Hangnest.

Two taken, but not eaten, by Harmonious Shrike-Thrush.

Two taken, but not eaten, by Cuban Mocking Bird,

Limax arborum.

Sept. 24,1910. Four eagerly eaten by Wall Lizards, which
wiped their mouths to remove the slime after swallowing them.

Milax sowerbyt.

Sept. 24,1910. One taken and pecked and wiped about in the
sand for a long time by Indian Dial Bird, which finally left it.

Another specimen was eagerly taken by Sulphury Tyrant, which
after pecking and crunching it in his beak, and banging it from
side to side against a ledge, exactly as Laughing and other
Kingfishers do, finally swallowed it whole.

ARACHNIDA.

OPILIONES (Long-legged Spiders or Harvestmen).

Phalangium sp. ?

Sept. 1910. One (immature) tasted but immediately rejected
by Pekin Robin; the same specimen then taken and eaten by hen
Scarlet Tanager.

One (immature) put into cage with several Curassows was tasted
in turn by specimens of Yarrell’s and the Globose, and ultimately
eaten by one of the Globose Curassows, when crushed beyond all
recognition.

I was led to suppose these Arachnida would prove on experiment
to be unpalatable owing to their possessing a pair of glands, one
on each side of the dorsal area of the carapace, which are known
to secrete an odorous fluid. As elsewhere recorded *, I have seen
a Mason Wasp, hunting Spiders, run down a specimen of Phal-
angiwm, but turn aside and let it go unhurt the moment he
touched it with his antenne. More experiments with birds and
lizards are required fully to substantiate my belief ; but the refusal
of the Pekin Robin to eat the Phalangiwm is very significant, and
it is quite evident that the Arachnid was not to the liking of the
Curassows.

* Journ, Linn. Soc., Zool, xxx. p. 268, 1909,

PALATABILITY OF SOME BRITISH INSECTS. 815

INSECTA.
Order LEPIDOPTERA.
Butterflies.

Group PIERIN#.
THe SmMaLt Wuire (Pieris rape).

July 31,1909. One male (dead) given to Capuchin (Cebus sp. a)
was taken at once, and eaten without being removed from the
mouth for inspection. This specimen, given with Huchloé carda-
mines (see p. 820), was used as a check upon the behaviour of the
monkey towards Huchelia jacobec and Melitea artemis (pp. 825
and 832).

Sept. 6, 1910. One offered to a Red-handed Marmoset was
inspected, but not touched; but was eagerly taken and eaten by
another animal of the same species. This Marmoset then ate a
specimen of Perarge megera, his behaviour suggesting that the
two butterflies were equally palatable to him.

May 26,1909. One chased at once by Shrike-Thrush and Dial
Bird, but evaded them and escaped through the partition into

next cage, where it was promptly caught on the wing by a
Fantailed Flycatcher and eaten.

One caught at once on wing by Great Tit and eaten.

Aug. 21 to 27, 1910. One greedily eaten by cock Silver
Pheasant.

One let loose in aviary skilfully dodged the swoop both of a
Shama and a Wood-Swallow, and escaped.

One given to Dial Bird, which took it from my hands and
damaged it by the peck so that it was unable to fly away. Again
and again he pecked the butterfly as it fluttered about on the
ground, but would not hold it. Ultimately it escaped under the
partition into the next aviary, where it was pounced upon by a
Weaver, which held it in his foot and ate it, leaving the wings.

Sept. 6,1910. One taken by Masked Wood-Swallow and eaten
after much delay and pecking. The bird evidently was not very
keen on the insect; but he would not allow any other bird to
take it from him. He did not once shake his head or wipe his
beak as if there was any distasteful flavour.

One female taken and eaten by Ludwig’s Bustard.

Sept. 7, 1910. One male and one female taken and eaten
eagerly and with equal avidity by Green Lizard.

Larva of the Small White (P. rape), fed on cabbage.
_ Sept. 21,1910. One tasted but rejected by Yarrell’s Curassow
and Globose Curassow.
THE GREEN-VEINED WHITE (Pieris napi).

July 31, 1909, One offered to White-tailed Mongoose, to three
06*

816 MR. R. I. POCOCK ON THE

Meerkats and to two Banded Mongooses. All rejected it after
smelling it except the second Banded Mongoose, which took
it with his paw, rubbed it in the sawdust, but would not
eat it.

N.B.—The forceps with which this butterfly was offered had
been previously used for Ocypus olens, Carabus violaceus, Ptero-
stichus niger and P. madidus, and some Timarche as well as
Coccinella, and probably the scent of these beetles was adhering to
the steel.

May 26,1909. One taken and eaten by Dial Bird, by Har-
monious Shrike-Thrush, and by Blue Rock-Thrush.

July 26, 1909. One male given to Silver Pheasant, was taken
from my fingers and swallowed instantly without being first
deposited on the ground.

One female given to same bird was treated in exactly the same
way.

These two I used as checks upon two specimens of Melanargia
galathea, both of which the Pheasant treated very differently,
spitting them out upon the ground after taking them from my
fingers, and pecking them about a great deal before swallowing
them (p. 827).

Aug. 21,1910. One male greedily eaten by Silver Pheasant.
This bird ate at the same time a specimen of Hpinephele jurtina,
showing an equal liking for both.

One male taken eagerly by Pekin Robin, which, after much
pecking and tasting, left the butterfly uneaten.

July 31, 1909. One male eaten at once by Brazilian Hangnest.

Sept. 20,1910. One left untouched by Fantailed Flycatcher.
Taken and tasted but left uneaten by Dial Bird. Taken by
Yellow-crowned Hangnest, which held the butterfly in his foot
against the perch, pecked off its wings and finally picked it to
pieces, and ate at all events most of it.

Note.—The Hangnests which ate these butterflies are much less
typically insectivorous in diet than the Flycatcher, the Pekin
Robin, and the Dial Bird, which refused them.

Tae Larce Waite (Pieris brassice).

Oct. 26, 1909. One taken from my hand and greedily eaten
by Lion Marmoset.

May 26, 1909. One taken at once by Syrian Bulbul and eaten ;
also by Harmonious Shrike-Thrush.

Oct. 26,1909. One taken from my hand and greedily eaten
by cock Silver Pheasant and by Honduras Turkey.

One taken by Shama and finally eaten, but not with any ap-
proach to the readiness with which he had just previously eaten
a Tortoise-shell and the #. jurtina. At one time I thought he
was going to give it up; but finally he swallowed it.

One liberated in aviary was chased up and down by three
Wood-Swallows which, however, owing to hesitancy at the moment

PALATABILITY OF SOME BRITISH iNSECTS. 817

of coming to close quarters, did not catch it. It escaped into
another compartment, and was promptly seized by the Harmonious
Shrike-Thrush, which ate it after a deal of pulling about and
tasting.

Aug. 21, 1910. Two males greedily eaten by cock Silver
Pheasant.

One male caught by Pekin Robin and eaten after some time,
the delay being caused not apparently by distastefulness, but by
the difficulty of getting rid of the wings which were left uneaten.
This bird held the insect to the perch with his foot when
pecking.

One male eagerly taken by Pearl-spotted Owl, which held it up
in one foot while pecking it. He pecked away for some time at
the thorax and wings without making much headway. He then
shifted it and pecked off the end of the abdomen. But as soon as
he got the flavour of the exposed tissues he shook his head and
repeated the shake with every taste, showing unmistakable signs
of disliking the flavour. Finally he hopped to another perch, put
the butterfly down, and after looking at it for a little time, flew
away. I thought he had given it up; but upon returning to the
cage ten minutes later the butterfly had disappeared.

One put into an aviary of Tanagers was chased by several birds
which, however, hesitated at the critical moment to catch it, as if
a little doubtful as to its nature. At last a male Scarlet Tanager
took it in his beak, but not having the instinct to use his foot to
hold it or to put it into a cranny, went on masticating it for at
least five minutes without showing any signs of dislike. He
apparently refrained from swallowing it on account of the wings.
Ultimately he was robbed by a female of the same species, which,
after getting rid of the wings, continued pecking and tasting and
shaking her head in the intervals, quite obviously not enjoying the
flavour. She managed the insect better than the male, jamming
it first into a split orange, and then between the leaves of a palm
to peck it the better. Ultimately she ate what was left of the
body.

One male offered to a hen King Bird of Paradise. She looked
at it and as soon as she saw the legs move took it, but dropped it
at once to the bottom of the cage. After careful and long in-
spection, she pecked it once or twice, but showed no eagerness to
eat it. Ithen gave the same insect to a Larger Hill Mynah,
which soon swallowed it, wings and all.

One male taken and eaten at once by Ludwig’s Bustard.

One male offered to Fantailed Flycatcher, but he would not
touch it. Taken and tasted by Dial Bird, but left uneaten. Also
taken and tasted by Black-winged Grackle, and left and sub-
sequently refused twice. Quickly eaten up by Harmonious Shrike-
Thrush.

Sept. 18 to 20, 1910. One caught on wing by Fantailed Fly-
catcher, which had just eaten a ‘Blue.’ He carried it to a
window-sill, but after one or two pecks left it. Once or twice the

818 MR. R. I. POCOCK ON THE

bird, after waiting a short while, tried it again, but finally left it
alone.

It was then taken by a Dial Bird, which, after pecking it about
for a short time, was robbed by the Sulphury Tyrant. The latter,
after tasting it, left it alone. I then gave the remainder of the
insect, consisting only of the thorax and wings, to a Yellow-
crowned Hangnest, which took it toa perch, and holding it in one
foot gradually pecked away the wings and dropped them, and
then pecked the thorax to pieces, eating little bits of it and
dropping others.

Pupa of the Lance Wurte (Pieris brassice).

Oct. 26, 1909. One offered to the Dial Bird which had fifteen
minutes previously eaten the larva, but he would not touch it.

Offered to Yellow-crowned Hangnest which had tasted and
dropped the larva. He looked at it but would not touch it.

Given to Harmonious Shrike-Thrush, which behaved just as
the Dial Bird had behaved with the larva, pecking it and dropping
it repeatedly to shake his head. He was then robbed of it by a
Common Mocking Bird, which, however, dropped it in the grass
from the perch, and made no attempt to recover it.

One offered to a Black-winged Grackle, a Javan Pied Mynah, a
Fantailed Flycatcher, and a Sulphury Tyrant, all of which tasted it
once, but not asecond time. A Common Mocking Bird persevered
a little longer, but finally dropped it and made no effort to pick
it up again. Given to Harmonious Shrike-Thrush, was eaten
without much hesitation.

Larva of the Lares Wurte (Pieris brassice).
Food not recorded.

Oct. 26, 1909. One taken by Yellow-crowned Hangnest, but
soon dropped. Pounced upon by Dial Bird, which after many
trials, pecking it and shaking his head after every taste, at last
swallowed it; but he was evidently very uneasy for some twenty
minutes afterwards, periodically shaking his head and opening his
mouth and straining as if trying to vomit something nauseous.

Larvee of the same fed on 7ropeolum (so-called Nasturtium).

Sept. 13, 1910. Three eaten readily by Silver Pheasant and
Reeves’s Pheasant.

A small one given to Pekin Robin, which obviously did not
like the flavour. He pecked it about in the sand for a long time,
vigorously shaking his head after each taste. Ultimately, how-
ever, he ate it. I then gave him asa test the larva of a Noctua
(see p. 835), which he also took and very soon swallowed entire
without once shaking his head or evincing any sign of dislike.
He then took a second and larger brassice-larva, treating it just
as he did the first, but tackled it with still greater reluctance,

PALATABILITY OF SOME BRITISH INSECTS. 819

allowing himself to be robbed of half of it by another bird of the
same species. The two finally finished it between them.

One given to a Shama, which after pecking and tasting it for
a long time, with much headshaking, left it. It was then tasted
by a Wood-Swallow, which left it after one peck. The Shama
then tried it again, but left it. ‘Then a Red-vented Bulbul took
it, but soon dropped it. The Shama then tried it again and
ended by eating it. This Shama was the same bird that ate the
Coccinella 7-punctata (p. 846).

One given to Kagu, which after several attempts left it; and
immediately afterwards greedily ate the larva of a Voctua (p. 835).
This same Kagu ate 7imarcha tenebricosa.

One taken by Green Hangnest, which at the time was greedily
eating mealworms. ‘The bird finally ate it, but evidently did not
much like it, putting it down several times, and wiping it in the
sand.

One given to Pearl-spotted Owl, which dropped it at once.

One given to Butcher Crow, which dropped it directly; but
afterwards picked it up and swallowed it whole. Immediately
afterwards, however, he vomited it up and left it on the bottom
of the cage.

One smelt, but not touched by Common Marmoset, and by
Capuchin.

One eagerly eaten by Meerkat.

Sept. 21, 1910. Larve of the same, fed on cabbage (Brassice).
Taken and eaten by :—

Elliot’s Pheasant, Reeves’s Pheasant, and Silver Pheasant.

Vulturine Guinea Fowl. Crested Guinea Fowl. Ludwig’s

Bustard. Vigors’s Bustard. S. American Thicknee.
Cariama. Crested Curassow. Nigerian Ground - Horn-
bill.

Also by Meerkats and Banded Mongoose.

Tasted but rejected by:—Shama, Red-vented Bulbul, Green
Hangnest, Black Hornbill, Elate Hornbill, Trumpeter, Yarrell’s
Curassow, Globose Curassow, Crested Curassow, and Red-
tailed Guan.

Notes.—The nature of the food of the larve did not appear to
affect their taste. The Green Hangnest, it is true, refused larve
fed on cabbage, having a week earlier eaten one fed on Z’ropeolum,
but the bird was not eager for the latter, and I do not think this
refusal of the former can be taken as strong evidence that he
found them more unpalatable than the others. It is interesting
that the Pheasants and Guinea Fowl, that is to say, Asiatic and
African Gallinaceous birds, ate the larvee eagerly, while the
S. American Curassows and Guans, with the exception of one
Crested Curassow, refused them after many trials, and much head-
shaking. One Curassow eagerly ate the larva of the Woctua
(p. 835) after refusing that of P. brassice.

820 MR. R. I. POCOCK ON THE

THE ORANGE-TIP (Huchloé cardamines).

July 31, 1909. One male given to Cebus (sp. «) was seized at
once and stuffed into his mouth. He took it out, looked at it,
smelt it, then ate it without hesitation.

This was a check experiment upon the behaviour of the monkey
towards Huchelia jacobee and Melitea artemis. He showed much
greater alacrity in eating the cardamines than either of the others.
A Pieris rape given at the same time he ate without removing
it from his mouth.

May 26, 1909. One male taken by the Harmonious Shrike-
Thrush after a few moments’ inspection and eaten entire, wings
and all, with much less delay in the way of pecking and scraping
on the soil than the same bird displayed when dealing with
M. artemis and A. euphrosyne. Tested by this bird, H. cardamines
appeared to be more palatable ; but it is possible, though I do not
think probable, that he ate it with less delay because he had just
previously been robbed of the specimen of Argynnis euphrosyne
by not swallowing it at once.

Group NyYMPHALINA,

THE SMALL ToRTOISE-SHELL (Vanessa urtice).

Oct. 26, 1909. One taken and eaten by Shama which had just
previously eaten Hpinephele jurtina.

Hoopoe, Black-winged Grackle, and Harmonious Shrike-Thrush
very eager to take one, but it was secured by the Grackle,
which, however, was robbed by the Shrike-Thrush, the latter
eating the butterfly in about half a minute without any signs of
dislike such as shaking his head or wiping his beak.

Sept. 7, 1910. One taken and greedily swallowed, wings and
all, by Dial Bird.

Sept. 18, 1910. One caught on wing by Fantailed Flycatcher,
who carried it to a perch, but after a few tastes and pecks
dropped it to the ground. Whether this was done intentionally
or accidentally I cannot say, but the bird made no attempt to
follow up the insect. I then gave it to a Dial Bird, which, after
pecking it for a short time, was driven off by a Sulphury Tyrant.
This bird, however, did not touch the butterfly. I then offered
it to a Bulbul and a Yellow-crowned Hangnest; but neither
touched it. I then offered it again to the Dial Bird, who finished
it, but with no show of appetite. I am unable to say whether
the indifference shown by the birds to this butterfly was due to
its being distasteful or to the experiment being made at 5 P.m.,
when the birds had been feeding off and on through the day.

Pupa of Vanessa urtice.

June 24, 1909. One placed on a branch near a Shama was
taken after a good deal of preliminary inspection but was soon

PALATABILITY OF SOME BRITISH INSECTS. 82)

flicked away and fell to the ground. The bird made no attempt
to recover it. I then again put it on the branch by his side, and
on this occasion he pecked at the little stem to which the pupa
was attached. A hen Black Tanager was the next to try it. She
broke the shell and getting the taste flew away with the pupa and,
T think, ate it. At all events she flew up to the top of some
brickwork where I could not see her clearly, and presently came
down again without the pupa; and on going upa ladder to look
for the pupa, I could find no trace of it.

One offered to Syrian Bulbul was taken after some scrutiny.
The bird flew away with it and pecked it, but seemed greatly
bothered and puzzled by the tightness with which it adhered to
the twig. He was unable to detach it from the twig, and finally
left it. I then offered it to a Fantailed Flycatcher ; but
could not induce this bird to touch it, although he scrutinised
it carefully and was hovering round me the while, apparently
remembering that on previous occasions I had given him butter-
flies. I then gave it to the Harmonious Shrike-Thrush, which
took it, pecked away at it until he broke off the tail-end and ate
it. He then pecked off another piece and ate it, showing no sign
of dislike. He then left the larger piece; but soon returned,
broke it up, and finally ate it piecemeal.

From watching the behaviour of these birds, I should say that
these pupe are unpalatable only to the extent afforded by the hard-
ness and toughness of the chitinous integument. The birds that
tasted them after breaking the exoskeleton, showed no signs of
disliking the flavours. Those that took them—and the Flycatcher
could not be induced even to attempt it—did so after scrutinising
them in a way that suggested doubt as to their belonging to the
category of eatable things. They did not appear to me to know
what they were; and none of the many insectivorous birds in the
aviary showed the least sign of eagerness when I first put the
pupa on a perch, waiting to see which would be the first to come
down. It was only when I placed it about a couple of inches
from the Shama, a tame and fearless bird, that he took it. The
Tanager came, and after her the Bulbul, when they had seen the
Shama’s attempt, or at all events after the Shama had first tackled
it. These birds are accustomed to visitors and keepers bringing
food into the aviary; and I think it probable that the Shama
was induced to peck at the pupa merely because it was definitely
offered to him.

I suspect that this pupa is protected in the first place by its
likeness to things inanimate, and in the second place by the
toughness of its integument which does not readily yield to a
peck, and is quite in keeping with the general impression of life-
lessness suggested by the colour, shape, and immobility of the
whole pupa. J may add that I did not see the pupze move when
pecked by birds, although they did so when handled by myself.

829 MR. R. I, POCOCK ON THE

Young larve of Vanessa wrtice.

June 24, 1909. One eaten without hesitation by Brazilian
Hangnest, and by Common Mocking Bird; two by Shama; two
by Orange-headed Ground-Thrush, and one by Harmonious
Shrike-Thrush.

Two taken and tasted but whisked away by Larger Hill Mynah.

One taken and tasted but dropped by North American Cat-bird,
which refused to touch a second.

~ One pecked and tasted many times, but finally rejected, by
Fantailed Flycatcher.

THE Peacock (Vanessa 20).

May 26, 1909. One fluttered to ground and rested with wings
closed. A Fantailed Flycatcher flew down to inspect and was
preparing to peck, when the butterfly opened its wings and moved
them slowly up and down. The transformation seemed to dis-
concert the bird, which made no attempt to peck, but danced
round the insect at a distance of about three inches. A Shama
and, another Flycatcher, which joined the first, behaved in the same
way. A Syrian Bulbul then flew down and drove the three away.
After inspecting the butterfly for about half a minute, he pecked
the ocellus of the anterior wing of the left side; the second peck
struck the ocellus of the anterior wing of the right side ; the third
the ocellus of the posterior wing of the left side, tearing a piece
out. He was then driven away by a Sun-Bittern, which looked
at the butterfly for some two minutes, but made no attempt to
peck it, although it excited his interest. I then removed the
Bittern ; and the Bulbul returned at once, seized the butterfly by
the head and thorax, flew away with it, and devoured it.

One fell to floor of aviary with wings closed, and was at once
seized by Syrian Bulbul, before its wings opened, and was carried
away and eaten. A second Bulbul of this species pursued the
tirst; but I do not know which of these two birds was the one
that ate the zo first introduced.

The two features of interest in the first experiment with this
species were, first, the manifest disconcertedness of the three birds
by the sudden display of colour and the slowly waving wings of
io (my wife, who was with me, said at once, ‘‘They are afraid
of its eyes”); and secondly, the consecutive pecking of three of
the ocelli by the Bulbul. It can hardly have been by accident
that the ocelli were accurately struck three times running.

Aug. 21, 1910. A specimen let loose in aviary was chased by
a number of Tanagers and other small birds and was caught by a
Scarlet Tanager. The latter, however, was robbed by a Pekin
Robin, which ate the insect without showing any signs of dislike,
the delay of five minutes in finishing it off being caused by the
difficulty of managing the wings which the bird ultimately broke
off and left uneaten.

PALATABILITY OF SOME BRITISH INSECTS. 823

THE Rep Apirat (Pyrameis atalanta).

Aug. 21, 1910. One taken and eaten greedily by Lion
Marmoset.

One pursued by Shama, which grabbed it by the hind wing and
thereby lost the butterfly, which flew away and escaped through
the wires of the aviary.

Tue Patnrep Lavy (Pyrameis cardui).

Aug. 27,1910. One given to Pearl-spotted Owl was taken at
once and swallowed entire after a little preliminary pecking.

This was a test experiment to ascertain the meaning of the
bird’s behaviour towards Pieris brassicw (see p. 817).

Araschnia levana. Late summer form prorsa.

July 8,1911. One given to Harmonious Shrike-Thrush, an
Australian bird, was taken at once, but after being pecked and
tasted for some little time, was rejected. The remains were then
greedily eaten by a Wood-Thrush, from North America. A fresh
specimen given to this same Wood-Thrush was just as readily
swallowed ; but the Shrike-Thrush upon taking another, treated
it as before, wiped it in the sand, shook his head, and allowed
himself to be robbed by a Black-chinned Laughing Thrush, which
ate it and another without hesitation.

One taken and eaten, but very slowly and with much pecking
about, by a Hoopoe, which, after swallowing the last particle,
appeared to try to vomit it back but without success.

A Blue Rock-Thrush and a Common Rock-Thrush, both
European birds, each ate one greedily.

One pecked and tasted for some little time by Orange-headed
Ground-Thrush, which obviously did not care for the favour, and
allowed himself to be robbed by the Blue Rock-Thrush mentioned
above.

One liberated in aviary dodged the pursuit of a Shama and a
Sibia with great skill, and escaped.

One given to Shama was pecked and tasted for some time, but
the bird allowed himself to be robbed by a Wood-Swallow, which,
after much pecking, swallowed the butterfly.

This performance was repeated exactly when one was given to
the Sibia, the same Wood-Swallow taking it from him; but I
think the Sibia would have eaten it ultimately.

One given to Grey-headed Friar Bird, from Australia, was
taken and tasted for a long time and then dropped, given again
to the same bird, was again tasted and dropped. The remains
were then eaten without much delay by a Larger Hill Mynah.

One given to a Dial Bird was taken and after much tasting
was resolutely rejected. The remains were then given to a

824 MR. R. I. POCOCK ON THE

Sun-Bittern, which persevered for some time but finally rejected
them.

The only birds which ate the butterflies quite readily were
the two species of Rock-Thrushes, the Wood-Thrush, and the
Black-chinned Laughing Thrush. To the others they were
obviously more or less distasteful, the most significant rejection
being by the Shrike-Thrush, which on previous occasions has
eaten almost every insect offered to him.

N.B.—These experiments were made between 4 and 5 P.M.,
when the birds had been feeding throughout the day.

July 9, 1911. One eaten readily by Black-headed Lemur,
one by Meerkat, two by Common Indian Mongoose.

Three eaten readily by two Wall Lizards.

Two eaten readily by Silver Pheasant, and one fairly readily
by Mantchurian Crossoptilon (Pheasant).

One given to White-eared Scops Owl was taken at once but
dropped as soon as tasted.

Experiment repeated with same result.

Experiment repeated with same result with another specimen
of the same species of Owl.

One given to Pekin Robin was taken at once, but put down
upon the ground. For fully five minutes the bird continued to
peck it and shake his head. He would neither eat it himself
nor allow the other birds to take it from him. Ultimately he
pecked it to pieces; but I cannot say whether he ate particles or
wasted them on the ground. One thing was quite clear. He did
not find the flavour to his liking.

Dark Green Frivitiary (Argynnis aglaia).

July 21, 1909. One let loose in aviary was chased by Black-
headed Sibia and Fantailed Flycatcher, but eluded both and
escaped into a crevice. This is the first butterfly I have seen
dodge the Flycatcher, which is extraordinarily adept at taking
insects on the wing. I then gave it to the Spectacled Thrush, and
he ate it after he had succeeded in shaking off its wings. The
bird was keen not to lose it, and drove away the Flycatcher
whenever he ventured near.

SILVER-WASHED F'RITILLARY (Argynnis (Dryas) paphia).
July 26,1909. One caught on wing and eaten with avidity
by Fantailed Flycatcher.
Also used as check upon Melanargia galathea which the
Flycatcher had rejected (see p. 827).
July 31, 1909. One eaten readily by Brazilian Hangnest.

PEARL-BORDERED FRITILLARY (Argynnis (Brenthis) ewphrosyne).

May 26 to 31,1909. One eaten by Silver Pheasant. For
details see under Melitea artemis (see p. 826).

PALATABILITY OF SOME BRITISH INSECTS. 825

Two specimens given respectively to Brazilian Hangnest, and to
Saturnine Mocking Bird, were eaten much more readily than
were specimens of J/. artenvis offered to the same birds (see under
Melitceea artemis).

One female taken by Harmonious Shrike-Thrush, but not eaten
readily. While this bird was pecking the butterfly and wiping it
on the gravel, he was robbed of it by a Red-vented Bulbul; the
latter was in turn robbed of half of it by a North American
Mocking Bird. The two finished it between them.

SMALL PEARL-BORDERED FRittLuary (Argynmnis (Brenthis) selene).

May 31,1909. One taken and eaten by Capuchin, but without
relish.

One taken and eaten by Capuchin (Cebus sp. c), with obvious
avidity.

GREASY FRITILLARY (Melitcea aurinia or artemis).

May 26 to 31, 1909. One male given to same specimen of
Cebus that took the Huchelia jacobee five minutes previously.
He behaved in exactly the same way towards it. Stuffed it into
his mouth, but the moment he got the flavour or the feel, took it
out in his hands, pulled it to pieces, cautiously tasted it, and then
ate it, but with no great show of satisfaction.

One taken and eaten by Capuchin (Cebus sp. 6), but with great
hesitation and no particular signs of relish. This monkey also
ate one Cenonympha pamphilus, one Argynnis selene, and one
Thanaos tages; but treated them all in the same way, evidently
not caring much for any of them. In this particular he showed a
marked contrast to the two other examples of Cebus, sp. a and c,
used for these experiments.

One male offered to Meerkat, taken and eaten at once. Kager
for more.

One male offered to Capuchin, taken and eaten at once. Eager
for more.

One male offered to White-handed Lemur, which after carefully
smelling it, refused it.

Same one offered to Crowned Lemurand White-fronted Lemur,
was smelt and refused in the same way.

Offered to Black Lemur, was smelt, then carefully taken into
the mouth, but was then pulled out with the hand; then again
tasted, but rejected as if distasteful, the tongue being rapidly
protruded and drawn back through the front teeth as if to scrape
off something unpleasant, perhaps scales.

One offered to Diana Monkey, was taken and eaten piecemeal,
apparently with relish.

The mammals above mentioned had not been fed, and were
without exception hungry.

826 MR. R. I. POCOCK ON THE

One taken by Brazilian Hangnest, which pecked at it, ate a few
pieces as if testing its flavour, then let it fall from the perch to the
ground, and left it there.

One taken by Saturnine Mocking Bird, which shook it about,
pecked it, ate a fragment or two, then left it.

One taken by Brazilian Hangnest, which pecked it several
times, and finally ate it. The Mocking Bird then returned, and
after many trials finished off the remains of the first specimen
that had been left by the Hangnest and of the second that had
been left by himself. It was quite evident that neither of these
birds found much satisfaction in eating these butterflies.

One female liberated in aviary, caught on wing by Garrulous
Honey-eater, and eaten without delay.

One female taken by Blue Rock-Thrush, but left on the ground
after being pecked. Suspecting that his leaving it was due to my
propinquity, | moved away and told the keeper to throw it to him,
He then caught it on the wing, and ate it. He then came close
to me on a perch and eagerly took another specimen (male) from
my hand, then a third (female), and ate both greedily.

Two given to Silver Pheasant were taken and eaten, but with a
great deal of pecking and tasting. Comparing this bird’s behaviour
towards them with his manner of eating Pieris napi and rape, |
am quite sure he found them to a certain extent unpalatable. I
thought at first that he merely disliked the wings. To test this
I gave him immediately afterwards a specimen of napi. He took
it from my hand and put it on the ground; then tasted it, and
without more ado swallowed it. I then gave him a specimen of
rape. He took it from me, and without putting it on the ground
ate it up. I then gave him a specimen of Perarge megera, which
flew into a bush. He went after it, found it, caught it with the
dexterity of a ‘practised hand,’ but treated it exactly as he
treated the artemis, pecking and whisking it about, ultimately
after much delay eating it piecemeal, but with what might be
described as a very dubious air. He behaved in a precisely
similar manner towards an example of Argyniis ewphrosyne.

Tam convinced that no one who had seen this Pheasant eat
these five butterflies, could have doubted for a single moment that
he found the ‘ Whites’ pleasant to taste, and the ‘ Fritillaries’ not
altogether to his liking.

One male offered to Larger Hill Mynah was taken and eaten,
but with no great relish, being frequently dropped and picked up
again, and scraped in the sand.

One male offered to Levaillant’s Barbet, which took it and be-
haved towards it in exactly the same way as the Mynah. The
birds appeared to dislike the wings, and to want to get rid of
them.

One male offered to Fantailed Flycatcher, which after a little
inspection pecked it and took it, but was robbed by a Syrian
Bulbul, which ate it.

Two males taken and eaten by Shama,

PALATABILITY OF SOME BRITISH INSECTS. 827

One male taken and eaten by Cape Robin-Chat.

One male taken and eaten by Indian Orange-headed Ground-
Thrush, after being pecked and rejected by Hoopoe.

One male taken and eaten by Harmonious Shrike-Thrush.

One female taken and eaten after a great deal of pecking and
delay by Indian Black-headed Sibia, which was chased for it by a
Syrian Bulbul.

One female taken and eaten, after a few moments’ inspection
and biting at the wings before the position of the body was found,
by a Sand Lizard. A Dugés’s Lizard came up while the butterfly
was being chewed, and after tasting it once or twice, attacked the
Sand Lizard to make him relinquish his hold.

One male taken by the same Sand Lizard after he had finished
the first specimen. I then made him drop it; and offered it toa
Wall Lizard, which took it without delay and swallowed it.

Group SATYRIN#.

THe MarsieD Waite (Jelanargia galathea).

July 24,1909. Asa check I first of all offered a specimen of
P. napi to the cock Silver Pheasant. He took it from my fingers,
and without hesitation swallowed it and turned eagerly for more.
I then gave him a galathea, which he just as eagerly took, but
promptly lowered his head to the ground and spat it out. He
persevered with it, however, and after a little pecking and shaking,
ate it. Ithen tried him with another napi. He took it and
swallowed it at once, not hesitating for a single moment, exactly
as he had done with the first one. Then I gave him another
galathea, which he took but immediately put out of his beak upon
the ground ; but after some pecking and tasting he swallowed it.

I consider this bird to have rather a refined taste for insects:
and I can now tell tolerably accurately by his behaviour whether
he likes one or not. And Iam quite sure that he found napi
very palatable and galathea not so.

I then let a galathea loose in the aviary, and it was promptly
caught on the wing by a Fantailed Flycatcher, which flew with
it to the ground, and after pecking, pulling and shaking it about
for a minute or so, gave it up and took no further notice of
it. As a check I then tried him with Aphantopus hyperanthus,
which he caught in the same way, and very quickly demolished.
I then gave him another galathea, which he caught and pecked
and shook for some little time; but he would not eat it. Asa
further check I gave him Argynnis paphia, which he caught and
disposed of as quickly as he had disposed of the hyperanthus.

I noticed that some of these galathea had darker spots below
than the others. Thinking that perhaps this might be a sexual
difference, I gave one of each kind to the Pheasant and to the
Flycatcher; but the birds behaved in exactly the same way
_towards them.

One caught and eaten by Dial Bird; also by Orange-headed

828 MR. R. I. POCOCK ON THE

Ground-Thrush, and by White-cheeked Bulbul. The latter was
robbed by the Harmonious Shrike-Thrush ; but recovered the
butterfly and ate it.

One given to Sulphury Tyrant, who pecked it and shook it for
a long time, then allowed the Shama to take it from him.
The Shama ate it. This was a galathea with lighter spots
below.

One given tothe same Shama was also eaten, This was a darker
spotted specimen.

The specimen above alluded to that was rejected by the
Flycatcher, was eaten by a North American Cat-bird (Thrush).

One offered to Australian Bustard, was taken from my hand
and swallowed at once.

One given to Meerkat, was taken and eaten without hesitation.

One given to Capuchin, which by his rejection of the Telephorid
peetle (p. 840) had shown himself to be more particular in taste
than some others of his species, was eaten, but by no means
greedily.

With the exception of the Australian Bustard none of the
birds that ate the galathea did so with great alacrity. Swallowing
them was in all cases preceded by a varying amount of flicking
and shaking and pecking. When I began my experiments I
thought this behaviour was due to a wish to get rid of the wings ;
but I am now doubtful about this, and believe that in many cases
at all events it indicates dislike of the taste. When a butterfly is
really to the liking of a bird, he disposes of the insect as fast as he
can, without paying much attention to the wings. This struck
me to-day particularly in the case of the Silver Pheasant when
eating the api, and of the Flycatcher when eating the hyper-
anthus and the paphia. The paphia especially was a large-winged
butterfly for so small a bird; and yet he swallowed it, wings and
all, in a few seconds.

Tue Meapow Brown (Lpinephele jurtina x janira).

July 21, 1909. One female eaten at once by Lion Marmoset,
which had previously refused to taste the malacoderm beetle
(Rhagonyche) and the Saw-fly (Allantus arenatus).

One female taken from my hand by Spectacled Thrush, but
made his escape. Caught on wing by Fantailed Flycatcher and
eaten at once.

One female eaten at once by Common Pheasant.

July 31, 1909. One male caught on wing by Black-headed
Sibia and eaten at once; another (female) caught on wing and
eaten without delay by Fantailed Flycatcher.

Aug. 21,1910. One male greedily eaten by Silver Pheasant.
This bird at the same time ate with equal avidity a male specimen
of Pieris napi. Her behaviour indicated no difference of taste
between the two butterflies.

Oct. 26, 1909. One female taken and eaten fairly readily by
Shama.

PALATABILITY OF SOME BRITISH INSECTS. 829

Tue Larce Hearn or GATEKEEPER (Hpinephele tithonus).

July 31, 1909. One eaten at once by Brazilian Hangnest.

Aug. 25, 1910. One caught and quickly swallowed entire by
Pekin Robin.

One male taken and swallowed entire with scarcely any delay
by Pearl. spotted Owl.

Sept. 20, 1910. One caught on wing and eaten without delay
by Fantailed Flycatcher. This bird would not touch two White
Butterflies (P. brassice and napi), offered one just before and one
just after it took tithonus.

THe Rinewer (Aphantopus hyperanthus).

July 26, 1909. One caught on wing and eaten with ail speed
by Fantailed Flycatcher.

I used this specimen as a check upon galathea, which the bird
had just refused to eat after catching it and pecking it about for
some time.

One gobbled up at once by Silver Pheasant.

Tue Smatt Hearu (Cenonympha pamphilus).

May 26,1909. ‘Two taken and eaten at once by Fantailed
Flycatcher.

One taken by Fantailed Flycatcher which was at once chased
by Syrian Bulbul.

One seized by Orange-headed Thrush, which after carrying it
about gave it through the bars of the partition to another bird of
the same species. This was deprived of it by a Bower Bird, which
earried it about, perhaps as a possible ornament, since he made
no attempt to eat it.

THe WALL Burrerrty (Perarge megera).

May 31,1909. One taken and eaten by Capuchin (Cebus sp. b),
but without apparent liking (see under Melitea artemis).

One taken and eaten with avidity by Capuchin (Cebus sp. c).

Aug, 25, 1910. Kagerly taken and eaten by Red-handed
Marmoset.

May 31, 1909. One eaten by Silver Pheasant (see also under
Melitea artemis).

Aug. 25,1910. Two (male and female) caught and greedily
eaten, wings and all, by Pekin Robin.

Sept. 5 to 7,1910. One female taken at once’ by Ludwig’s
Bustard, which, however, let it escape. It was caught on the
wing and quickly eaten by a Larger Hill Mynah. Two more
specimens (female) eaten by Black-winged Grackle and by
Dial Bird.

Proc. Zoou, Soc.—1911, No. LVTI. 57

830 MR. R. I. POCOCK ON THE

\
THE GRravuine (Satyrus semele).

July 31, 1909. One caught on wing by Fantailed Flycatcher
and eaten with all speed.

Group Lyc#NID&.

Common Buus (Lycena icarus).

Oct. 26, 1909. One given to Shama, but it avoided him and flew
through into the next compartment, where it was captured smartly
by a White-browed Wood-Swallow, and eaten as soon as the bird
could get peace from the pursuit of two other Wood-Swallows in
the same compartment.

N.B.—These Wood-Swallows were desperately keen to get
the butterflies with which they saw me feeding the Shama in the
next compartment.

One male let loose in aviary containing Fantailed Flycatcher,
which I should describe as an expert butterfly-catcher. But the
Blue dodged him again and again, and got through into the next
compartment. Here again it avoided the swoop of one or two
birds whose identity I did not detect in my intentness in keeping
my eye on the butterfly. The latter then passed through to a
third compartment and settled on some yellow painted boarding,
which it did not match, and on which it was caught by a Brazilian
Hangnest, and quickly eaten.

One male caught deftly by Masked Wood-Swallow, which after
prolonged pecking and tasting, swallowed the body, having got
rid of the wings. This bird used its foot to hold the insect down.

One male caught by Pekin Robin and ultimately swallowed
entire; but the bird put the insect down many times before
swallowing it.

The behaviour of these two birds suggested that this ‘ Blue’
was not very palatable. Its size offered no obstacle to its being
swallowed at once ; but both birds delayed over the meal.

Two (male and female) given in succession to Pearl-spotted Owl
were taken and swallowed entire without delay.

Two (male and female) let loose in aviary were captured and
quickly swallowed entire by Pekin Robin.

Troe Brown Arcus (Lycena astrarche).

Sept. 18, 1910. One let loose in aviary was taken by a Cayenne
Tanager, which was quickly robbed by a Pekin Robin. The latter
ultimately swallowed it entire, after putting it down several times
before finishing it off.

One caught on wing and eaten at once by Fantailed Flycatcher.

One caught and eaten at once by Dial Bird.

THE SMALL Copper (Chrysophanus phleas).

Aug. 25,1910. One caught and swallowed quickly, wings and
all, by Pekin Robin.

PALATABILITY OF SOME BRITISH INSECTS. 831

Group NEMEOBIIN&.

THE Duke or Burcunpy (Vemeobius lucina).

June 15,1909. One given to Brazilian Hangnest was taken
and swallowed without any hesitation.

One given to Saturnine Mocking Bird was taken at once, but
not eaten eagerly. While she was pecking it about a Lesser Hill
Mynah flew up and took it away, but was in turn deprived of it
by the Mocking Bird, which then swallowed it quickly.

Two given to Silver Pheasant, which ate them with the same
eagerness as it had previously shown when tried with the
‘Whites.’

Group HESPERIIDA,

Tue Dryey Sxiprer (7hanaos tages).

May 31, 1909. One taken and greedily eaten by Dent’s
Monkey.

One taken and eaten cautiously by Capuchin (Cebus sp. 6). See
under J/. artemis.

THe Larce SKIPPER (Argiades sylvanus).

July 21 to 31, 1909. One caught and eaten at once by
Fantailed Flycatcher, and one by Brazilian Hangnest.

Moths.

Larva of Goat-Moru (Cossus ligniperda).

Sept. 20,1909. One taken first of all by Dial Bird, which after
a short time was driven off by Sulphury Tyrant. Both, after
pecking it, left it practically uninjured on the ground. It was
then taken by the Harmonious Shrike-Thrush. He kept it for
some time, pecking it about and was eager to prevent other birds
getting it; but was finally beaten in a ‘tug-of-war’ for it by a
Common Hangnest, which carried the grub to a bush, held it
against a branch with his foot, and pecked away for five minutes,
then voluntarily dropped it. It was then taken by a Dial Bird,
which persevered for a long time, pausing frequently between the
pecks, opening and shutting and wiping his beak. He was then
deprived of it by a Black-chinned Laughing Thrush, which kept
it for ten minutes, pecking and whisking it about without making
any visible impression on the skin. The head, however, was by
this time gone, and the bird pulled some soft tissues out of the
end and ate them. I then gave it to a Green Hangnest, but
after tasting it he let it drop and took no further notice of it.
I then gave it toa Leach’s Laughing Kingfisher, which after a
little delay swallowed it whole.

ihe

832 MR. R. I. POCOCK ON THE

Larva of the Lackny (Clisiccampa neustria).

Sept. 20, 1909. Seized at once by a Shama which flew away
with it, holding it by the head; but while he was adjusting it
for eating, the female Black Tanager grabbed the other end and
being victorious in the tug that ensued, carried away the larva
and, after a good deal of pecking, ate it.

Srx-sporteD Burnet (Anthrocera (Zygena) filipendule).

July 31, 1909. One placed on a branch was immediately seized,
but flicked away by a Black-headed Sibia, which made no attempt
to follow it up but flew away, shook his head once or twice, and
wiped his beak.

Sulphury Tyrant then pecked it and flicked it away; and tried
it again with the same result, and left it.

Harmonious Shrike-Thrush took it eagerly, wiped it on the
ground several times, then jammed it into a forked branch and
started gingerly pulling it to pieces with much shaking of his
head and wiping of his beak. He then broke it in two pieces;
flew away with one and pushed it into a cranny and still per-
severed. He then broke another piece off, and stuck it in a cleft
branch ; but finally left it. I did not see him eat any of the
moth although he may have swallowed small particles. In any
case there were pieces of it left in the places where he had
fixed them.

Tue Crnnapar Morn (Luchelia jacobee).

July 31, 1909. One given to Meerkat, which caught it on the
wing with a snap, devoured it with every sign of relish, and
seemed eager for more.

One given to Capuchin (Cebus sp. a), which stuffed it into his
mouth at once, chewed it, then hastily took it out again, ap-
parently finding he had something either unusual or unpleasant
on his tongue; smelt it, pulled it to pieces with his hands, and
finally ate it, but with a good deal of doubt as if undecided as to
whether it was nice or nasty.

July 5, 1909. One specimen offered to a Fantailed Flycatcher
was immediately seized and pecked and tasted, and then rejected.
The Shama then tried it, and treated it in the same way, finally
rejecting it. A second Flycatcher then tasted it, and rejected it.

Another specimen of the moth let loose in this aviary flew
through the wires into another compartment, and was captured
on the wing by a Pied Grallina. He pecked it once or twice, and
tasted it, then flew away. A Cuban Mocking Thrush then came
up, and while he was looking at it and hesitating to peck, the
Grallina came back, drove away the Mocking Bird, seized
the moth and gradually ate it, holding it in one foot and
pecking it to pieces,

PALAVABILITY OF SOME BRITISH INSECTS, 833

Larva of Cinnabar Moth.

Aug. 15, 1909. Inspected but not touched by English
Thrush.

Offered to many fowls, only one of which pecked it, but dropped
it at once and took no further notice.

Maepie Mors (Abraxas grossulariata).

Aug. 1909. Offered to fowls, was inspected by several, but
only pecked by one, which at once dropped it, and made no
further attempt.

Small Green Geometra larva, probably of Cabera
pusaria or exanthemaria.

May 26, 1909. One taken without any hesitation by a Shama ;
but dropped. Then taken a second time, and dropped. When
preparing to take ita third time, he was deprived of it bya
Black-headed Sibia, which after spending a few seconds adjusting
it in his beak, swallowed it. It appeared to me that the Shama
dropped this larva accidentally, owing to lack of skill in adjusting
it in his beak, rather than intentionally. He was just as eager
to take it, although dead, the third time, as the first.

THE SWALLOW PRoMINENT (Pheasia dicta or tremule).

July 12,1909. Flattened itself to the ground but was at once
pounced upon by the same Flycatcher that had just eaten the
Hemerobiid (see p. 835). The bird, without any hesitation, ate
it with all speed, being merely delayed by the trouble of adjusting
the wings. Both this moth and Mamestra persicarie betrayed
their identity as Lepidoptera by flying out of the boxes to the
ground, so I had no chance of judging whether the Flycatcher
or other birds would have been deceived by their procryptic
coloration.

THe Bure-tie (Phalera bucephala) (amago).

July 5, 1909. Not being aware of this moth’s propensity, I
picked it up by the wings, whereupon it immediately twisted its
abdomen round and ejected a stream of white fluid over my
fingers. | regret that I missed seeing this defensive device
practised on a bird. However, I placed the moth on a wooden
branch, and a Fantailed Flycatcher flew down to inspect it ; after
loooking at 1t for a few seconds, he flewaway. I then put it near
a Shama, who hopped up to it and almost immediately picked
it up by the thorax. The other birds in the aviary now became
interested and pursued the Shama, giving him no chance of
eating it. When on the wing he dropped the moth, and the
Fantailed Flycatcher, which had previously taken no notice of it,

834 MR. R. I. POCOCK ON THE

immediately pounced on it, and after one or two efforts swallowed
it at a gulp. :

My impression is that the Flycatcher did not suspect the moth
of being eatable until he saw the Shama take it. It certainly
looked very like an inanimate excrescence as it rested on the perch.
Presumably the moth had exhausted its intestinal artillery upon
me, because it shot out no more when seized by the birds, but
kept perfectly quiet without even flapping its wings, although the
Shama did not crush it, and having it end on by the thorax with
the moth’s head in his mouth, left the wings perfectly free to flap,
and the abdomen to wriggle, had the moth been disposed to
struggle. This behaviour, I take it, was a manifestation of the
deeply implanted instinct to keep absolutely still (commonly called
‘death-feigning’), which is so highly developed in many animals
with procryptic shape and colour.

THe Dor (Mamestra persicaric).

July 12, 1909. One flattened itself to the ground, and was
seized by the Fantailed Flycatcher that had eaten Pheasia tremule,
and was eaten with avidity, delay, however, being caused by the
bird’s desire to get rid of the wings as well as by being disturbed
by another Flycatcher and a Syrian Bulbul, which tried to de-
prive him of the moth. The Bulbul subsequently picked up the
pieces of wing and ate them.

Larve of the Briaut-LtivE Brown-rve (Mamestra oleracea).

Oct. 26, 1909. One given to Harmonious Shrike-Thrush was
taken after a moment’s scrutiny. He pecked it, and tasted it
three or four times, then swallowed it readily enough. His be-
haviour suggested to me a certain amount of caution at first, as if
he remembered the distastefulness of the pupa of Pieris brassice
which he had just previously eaten. The green hue of both gave
them a superficial similarity to one another. Having eaten the
one specimen of oleracea he was very keen to get the second.

This I gave to the Black-winged Grackle which a few minutes
previously had unhesitatingly left the pupa of P. brassice after
one taste. He took it, and after a taste or two proceeded to eat
it with avidity, not giving the Shrike-Thrush, who was hovering
near and following him up for an opportunity to snatch it, a
chance to do so.

Larva of Drinker (Cosmotricha potatoria).

May 26,1909. One thrown to floor of aviary, was followed
by many birds and secured by a female Black Tanager, which
earried it to a perch and proceeded to peck it and shake it for
about one minute. She then dropped it, and it was seized by
the Black-headed Sibia, but was dropped at once. The Tanager
thereupon tried it again; and again let it fall, this time almost

PALATABILITY OF SOME BRITISH INSECTS. 835

immediately. Two Fantailed Flycatchers then came up and in-
spected it. One of them pecked it, but let it alone after one
experimental taste. The larva was by this time dead. Thena
Sulpbury Tyrant came up, picked it up and after a peck or two
swallowed it.

Larva of Voctua (unidentified).

May 26, 1909. Two (fed on Tropeolwm, so-called Nasturtium)
eagerly eaten by Pekin Robin and by Kagu.

One (fed on cabbage) was readily eaten by Yarrell’s Curassow,
which had just rejected the larva of the Large White (Preris
brassice) and of the Small White (P. rape).

Order NEUROPTERA.

Hemeprosip (unidentified).

July 12, 1909. One turned loose in aviary was at once caught
on the wing by Fantailed Flycatcher and eaten without hesitation.
The bird wiped its beak two or three times ona branch afterwards ;
but I do not think this action can be regarded as a certain sign
that it wished to remove something unpleasant. It suggests the
possibility, however, especially in view of the fact that the action
was not repeated by the same bird after greedily eating Pheasra
dictea and Mamestra persicarie.

Large Black and Yellow Dragon Fiy
(Cordulegaster annulatus) male.

July 26, 1909. One pounced upon and eaten after a time by
Harmonious Shrike-Thrush.

Order ORTHOPTERA.
Common GRASSHOPPER (Stenobothrus sp.).

Sept. 6, 1910. One given to Pekin Robin was eagerly taken
and eaten, but not with great rapidity, the bird putting it on the
ground between the pecks, but without once shaking his head or
showing any signs of disliking the taste. He appeared to me to
be troubled by the insect’s legs.

Great GREEN GRAssHopPER (Locusta viridissima).

All the birds in the aviary were keen to get it. It was tackled
at once by a Dial Bird; but he was driven off by a Black Tanager,
who flew away with the insect and pulled it to pieces on the top
of a wall.

SumMATRAN Stick Insect (Lonchodes sp.).

Taken and eaten at once by:—Pinché Marmoset, Lion Mar-
moset, Douracouli, Capuchin, and Banded Mongoose.

836 MR. R. I, POCOCK ON THE

Taken in the hand, but put down untasted and unhurt by
Grey Lemur.

Taken and eaten at once by:—Silver Pheasant, Cartagenian
Motmot, Fantailed Flycatcher, two Dial Birds, Shama, Black-
chinned Laughing Thrush, two White-crested Jay-Thrushes,
Black-winged Grackle, Chinese Mynah, Brazilian Hangnest, and
Shrike.

The Shrike was too shy to take the specimen from my fingers,
so I threw it towards him on the sand, not seeing exactly where
it fell. He, however, saw the direction of the falling insect, and
hopped towards it, but somewhat to my surprise—for birds
seldom lose sight of thrown food—did not pick it up but looked
as if inquiringly up at me. After a little search I found the
small Stick Insect on the sand lying still with legs extended, and
looking exactly like a blade of green grass. When I stirred it
up and made it crawl, the Shrike was on to it in a moment; and
I have no doubt that he missed it in the first instance owing to
its resemblance to the grass blade.

I observed that several of the birds looked inquiringly, as I
should describe it, at the Stick Insects before taking them. One
in particular, the Harmonious Shrike-Thrush, usually one of the
keenest insect-eaters in the Gardens, hesitated on two occasions
so long before making up his mind to touch them that he was
promptly robbed of his prey, once by a Dial Bird and once by
the Black-chinned Laughing Thrush.

Two birds took them directly, but instead of eating them,
hopped about with them in their beaks. One of these, the Green
Hangnest, was deprived of his by a Chinese Mynah, which took
it from him through the partition bars of the next aviary; the
other, a Grey Struthidea, was similarly robbed by a White-crested
Jay-Thrush after a Collared Jay-Thrush had made several
attempts to get it from him.

Order COLEOPTERA.

Group GEODEPHAGA.

The species of this group used for the test belonged to the
Carabide, a family of carnivorous ground-beetles with an
exceedingly hard exoskeleton. Carabus violaceus is black with
blue reflections; the species of Péerostichus are dead black and
shine like pitch. Harpalus has pubescent elytra and is a little
less conspicuous.

Carabus violaceus.

July 51, 1909. One rejected, after being smelt by three Meer-
kats, two Banded Mongooses, and one White-tailed Mongoose.
The latter behaved towards it exactly as he did towards the
Ocypus oles (see p. 838).

PALATABILITY OF SOME BRITISH INSECTS. 837

Offered same specimen to Harmonious Shrike-Thrush, which
seized it eagerly but was robbed by the Dial Bird. I am sure by
the way they tackled the beetle that either of these birds would
have eaten it ; but the Spotted Bower Bird robbed the Dial Bird,
as in the case of O. olens, and finally finished it.

July 31, 1909. One dropped on to floor of cage of the
Meerkat which had just eaten a Zimarcha tenebricosa. He
pounced on it, but would not seize it as he did the Zimarcha. 1
think he bit it, but am not sure. However, by the way he
pawed it about I am convinced he did not care for it. While
he was holding and smelling it, he quite suddenly let it go and
vomited up the Zimarcha (see p. 841). The Carabus escaped
unhurt. I then gave it to a Capuchin which seized it, and was
proceeding apparently to eat it when another snatched it from
him and ate it without showing any marked signs of dislike, but
with no great avidity.

This species, like others of the genus Carabus, discharges from
its mouth when handled a most repulsive smelling fluid.

Mr. Beddard found that Lacerta ocellata ate this beetle.

Pterostichus (Abrax) striola.

July 26,1909. One taken by Sulphury Tyrant which shook it
and pecked it for some time until robbed of it by Spectacled
Thrush. This bird also pecked it and banged it about until
robbed by female Black Tanager, which ultimately ate it after
much pecking and tasting.

The delay in eating this beetle on the part of the birds that
tried it may have been due to its hard exoskeleton or to partial
unpalatableness from other causes. The hardness alone would, I
think, account for it.

One (dead) given to Silver Pheasant was swallowed entire with
very little delay. The bird, however, after taking the insect
from my fingers, put it on the ground as is his custom with
anything hard or with soft butterflies not quite to his liking.

July 31, 1909. One seized and bolted at once by Silver
Pheasant in exactly the same way that he had bolted the other
Carabide.

Pterostichus niger.

July 31, 1909. One smelt but rejected by three Meerkats;
snatched from the forceps by a Common Indian Mongoose,
which followed it and watched it, and smelt it as a cat does a
cockroach, but did not eat it, so I took it from the cage un-
injured. White-tailed Mongoose turned from it in disgust.

One seized and bolted at once by Silver Pheasant.

According to Mr. Beddard this beetle was eaten without
hesitation by Lacerta vivipara and another lizard: and with some
hesitation by Finches.

838 MR. R. I. POCOCK ON THE

Pierostichus (Steropus) madidus.

July 31, 1909. One smelt and refused by three Meerkats.
Seized and eaten by White-tailed Mongoose. This Mongoose is
a large animal approaching a cat in size.

One seized and bolted by Silver Pheasant.

One pecked twice by Elliot’s Pheasant, but escaped into the
grass unhurt.

One seized and eaten by Black-headed Sibia.

Harpalus ruficornis.

July 21, 1909. One pecked at twice by Silver Pheasant but
not eaten, the bird taking no further interest in it after the
second peck. The beetle escaped unhurt.

Group BRACHYELYTRA.

Devit’s CoacH-HorRsE or Cock-TaIL BEETLE (Ocypus olens).
(Uniformly velvety black in colour.)

July 31, 1909. One smelt and rejected at once by three
Meerkats, one Mongoose, one Banded Mongoose, and by the
White-tailed Mongoose that had just before eaten the Timarcha
(see p. 842). This Mongoose started away from the scent in a
way that reminded me of the behaviour of a person who finds a
bottle of smelling salts unexpectedly pungent.

Offered the same specimen to Harmonious Shrike-Thrush,
which tackled it at once, but while pulling it to pieces was
robbed by the Dial Bird, and this bird in turn was robbed by a
Spotted Bower Bird, which ate it.

Note.—The difference between the Viverrine mammals and the
birds in their behaviour towards the Ground Beetles (Carabide
and Ocypus olens) was very marked, and is to be in a measure
explained, I think, by the wide difference in their powers of
smell. ‘The beetles appear to be relished by the birds; but to be
nauseous to the mammalia, This perhaps is natural; because
the Passerine birds would seldom come across the Ground Beetles,
which are cryptozoic and largely nocturnal. The mammals like
the Meerkats, and the Mongooses, on the other hand, must
commonly find them as they grub about and hunt for food on
the ground. Therefore one would expect protective attributes,
if existing at all in these beetles, to be of a kind to guard them
against being eaten by Meerkats or insectivorous mammals of
similar habits.

The Silver Pheasant which ate these beetles is essentially a
diurnal feeder and would seldom find nocturnal beetles. After
seeing him eat the Péerostichi as if they were large seeds, I do
not understand why he did not eat: the Harpalus ruficornis
offered to him some time previously *.

* Mr. G. A. K. Marshall suggested at the meeting when this paper was read that
the Harpalus had retained while the Péerostichi had discharged their acrid juices.

PALATABILITY OF SOME BRITISH INSECTS. 839

Group LAMELLICORNIA.

Dune BEETLE (Geotrupes vernalis).

July 23, 1909. Offered to Pearl-spotted Owl, which blinked at
it, but refused to touch it. Offered to a White-eared Scops Owl,
was at once taken and held up in one foot ; but after a few pecks,
which removed some legs, it was let fall, no effort being made to
recapture it. Given to a Ludwig’s Bustard, was eagerly taken,
and swallowed whole after a few pecks.

The large CockcHaFER (JJelolontha vulgaris).

July 23, 1909. One dropped on floor of aviary was pounced
upon by Indian Dial Bird which had just before been trying the
Timarcha. He pecked it, hammered it with his bill, and after a
great deal of difficulty broke it in half. He evidently liked it,
because he would not give any other bird a chance of getting it.
However, when he had broken it up, the Harmonious Shrike-
Thrush secured one half and carried it away, and after pecking it
for a few minutes swallowed it. The Dial Bird in the meantime
finished off his portion.

Srac BEEerLe (Lucanus cervus) male.

July 31,1909. This I showed to some Capuchins, which evinced
the greatest eagerness to secure it, but no sign of fear. I gave it
to one, and his first act was to bite off the mandibles. This may
have been an accident, but it reminded me of the alleged action
of baboons in removing the stings of scorpions before they can do
any damage with them. He then bit off the legs, finding they
worried him, and sitting down munched up the beetle as if it had
been a bit of apple. On a previous occasion I gave a dead Stag
Beetle (male) to some Brush Turkeys. One seized it and was
promptly chased round and round the enclosure by the others,
which evinced the greatest keenness forashare. I could not wait
to see what ultimately happened to the insect.

Group LONGICORNTA.

Strangalia armata, the only species of this group experimented
with, is a black and yellow, somewhat wasp-like flower-haunting
diurnal beetle, with a very hard exoskeleton.

July 21 to 31,1909. One taken at once from my hand by
Silver Pheasant and eaten after a good deal of pecking and
breaking up. The way the bird persevered with this hard-shelled
beetle shows that his rejection of the Harpalus was not due to
its hardness (p. 838).

One offered to Fantailed Flycatcher, which, however, would
not touch it. Black-headed Sibia took it without hesitation, and
flying away with it pecked it to pieces and finally ate it. Further

840 MR. R. I. POCOCK ON THE

evidence of the bird liking the insect was shown by the way he
flew away with it when chased.

One taken and eaten by Dial Bird, which was apparently only
delayed in disposing of it by the hardness of the exoskeleton.

One taken and similarly disposed of by Great Barbet.

One eaten after being broken up and crushed by Brazilian
Hangnest.

Group MALACODERMATA,

The beetle of this group used for the experiment is a flying
diurnal flower-haunting species, with a soft exoskeleton. It is
quite fearless of exposure. Beetles allied to it commonly form
centres of mimetic attraction in the tropics.

TELEPHORID ( ? Rhagonyche fulva).

July 21,1909. Four offered to four Capuchins were eaten,
two readily and without examination, two after a good deal of
tasting and examination between the tastes.

Two offered to two Capuchins were taken into the mouth,
tasted, then taken out, wiped on the bars and left.

One refused by Ceylonese Macaque after being smelt.

One eaten by Mona Monkey after a good deal of tasting,
smelling and pulling about. This Mona also ate the bug 7’ropi-
coris rujipes (p. 847).

One offered to Lion Marmoset was taken in the hand, smelt,
and promptly dropped. The Marmoset then descended from the
perch, picked it up again, smelt it and dropped it. The beetle
crawled away unhurt.

One smelt once or twice by Meerkat, was rejected without
being tasted.

One taken by Silver Pheasant, was pecked twice and left alone.
Another offered to same bird was pecked once and left. One
taken by Fantailed Flycatcher was pecked and tasted, then left.
The same specimen was then pecked once or twice by a Shama
and rejected. Black-headed Sibia then tried it, but gave it up
and vigorously wiped his beak after a taste or two. Afterwards
he made another attempt with the like result. The Black
Tanager then took it, tasted it, wiped his beak and rejected it.

One caught on wing by Harmonious Shrike-Thrush was eaten
after much pecking and pulling about. Another was treated in
the same way by this bird.

Two specimens, one of which was dead, offered to and eaten by
Dial Bird.

One tasted two or three times by Shama but rejected.

One pecked by Black-chinned Laughing Thrush, but flicked
away. Pounced on and eaten by Dial Bird.

Although eaten by the Dial Bird and the Shrike-Thrush,
which ate most of the insects offered to them, and by some of the

PALATABILITY OF SOME BRITISH INSECTS. 841

Monkeys, there can be no doubt that this soft-shelled beetle
possesses distasteful attributes. Its rejection by the Meerkat,
which ate nearly all the insects offered to it with the exception of
Coccinella 7-punctata, was very significant, and suggestive of nasty
smelling secretions.

Group PHYTOPHAGA.

The three species of this group that were tested are well-known
species. They are slow-moving diurnal forms found on plants of
different kinds. ‘They are squat in shape, dorsally convex, and
have a very hard exoskeleton, the Ladybird (Coccinella) being in
addition exceedingly slippery and difficult to hold. The coloration
of the latter is orange with black spots. The others are uniformly
black or blue. TVimarcha tenebricosa, the familiar ‘ bloody-nose
beetle,’ is further notorious for the discharge from its mouth of
a crimson liquid, whence the trivial name is derived.

Chrysomela polita.

July 31, 1909. One oftered to Meerkat was smelt and refused.
Another Meerkat in the same cage took it in his mouth, but spat
it out; both then sniffed it as it lay on the ground, but would
not touch it.

The same specimen, offered to a Grison, was sniffed but not
touched. Snapped up by McCarthy’s Mongoose; but was at once
spat out and left. It was then taken and eaten by a Banded
Mongoose.

Query: Had the previous tasters exhausted the Beetle’s supply
of nauseous juices ?

One given to Dent’s Monkey was taken, rubbed between
the hands and in the sawdust, smelt, tasted, pulled about and
rejected. Picked up by Mona in the same cage, but rejected
after one taste. ‘This Mona had just eaten a living Bombus.

One given to Harmonious Shrike-Thrush was taken, pecked
and tasted for a little, then left. Picked up by Black-chinned
Laughing Thrush, was pulled to pieces, and rejected. This bird
may have eaten pieces of the beetle, but the other débris was left
on the turf. He did not appear to find it very unpalatable.
Possibly in this case the nauseous juices had been exhausted by
the Shrike-Thrush.

One pecked off a perch by Fantailed Flycatcher, but not
followed up. Pecked and tasted by Sulphury Tyrant, but left.
Then tried by Sun-Bittern, but also left, crushed but with
nothing missing.

Timarcha tenebricosa (=levigata).

July 23-31, 1909. One offered to a Meerkat was eagerly seized,
chewed up and swallowed without much hesitation. But while
this Meerkat was just afterwards occupied with the Carabus viola-
ceus (cf. supra, p. 837), he yomited the Timarcha, I do not know

842, MR. R. I. POCOCK ON THE

whether the sickness was caused by the smell of the Carabus,
which to me is nauseating, or to its taste, or by the irritation of
the stomach caused by the Zimarcha. I suspect the latter,
because the Meerkat refused to touch a second Zimarcha that
was offered to him.

One smelt and rejected untasted by two more Meerkats ;
taken by a third in the same cage, rubbed in the sawdust, but
left apparently uninjured.

One grabbed at once and eaten by White-tailed Mongoose,
which immediately afterwards heaved and went through the
action of vomiting without, however, ejecting the beetle. <A
second specimen was smelt and rejected with every show of
disgust by the same animal, which persistently refused for
the next two hours every beetle that was offered him, although
before eating the Zimarcha he had devoured a Pterostichus
madidus. One rejected without being closely smelt by a Banded
Mongoose which had eaten a Coccinella 7-punctata. Seized by a
second Banded Mongoose, and eaten after a good deal of rubbing
in the sawdust.

One offered to a Capuchin, one of the specimens which had
refused the Telephorid (hagonyche fulva) (p. 840), was taken,
smelt, and rejected.

One offered to another Capuchin was ultimately eaten piece-
meal, but with so much delay caused by handling, licking, and
inspection, that I am sure it was no great treat to him, especially
as he had every reason to eat it speedily because a bigger
Capuchin in the same cage, which had snatched the Carabus
from his grasp, was almost continually after him to get the
Timarcha. When monkeys like their food they gobble it up if
there is the least likelihood of another taking it.

One offered to a Vervet Monkey was accepted, pulled to pieces
and eaten, the exoskeleton being dropped to the ground.

This specimen of Zimarcha had been previously offered to a
Baboon (Papio sphinx) ; but he would not even touch it.

One put on the floor of aviary was pounced upon by Dial
Bird, which after continued pecking and hammering could
make nothing of it beyond breaking it in half at the waist.
Ultimately he left it. An Orange-headed Thrush then tried
the abdomen, but was driven off by a Hoopoe, which after
pecking and hammering it, gave it up. The Thrush then tried
again, and also gave it up. A Black-chinned Laughing Thrush
then had a turn ; but with the same result.

One given to Harmonious Shrike-Thrush which had eaten the
Coccinella. He persevered for a long time, but could not manage
it and flew away, leaving the beetle apparently unhurt. After
about five minutes the bird came back and tried again, this time
pecking off the legs and antenne of the beetle; but he would
not eat the body, and at last flew away and returned no more.

Sept. 18, 1910. One female taken by Kagu, well crushed,
then swallowed at a gulp. °

PALATABILITY OF SOME BRITISH INSECTS, 843

One male taken by Vigors’s Bustard, crushed and put down
with a head-shake ; then tasted by two Ludwig’s Bustards, the
three birds having alternate pecks at it, the Vigors’s Bustard
finally swallowing it.

One female well tasted, but rejected by Wood-Swallow, Black-
winged Grackle, Javan Pied Mynah, and Black-chinned Laughing
Thrush: also by Sun-Bittern, which persevered for a long time,
repeatedly washing the beetle in the water-trough, and taking a
drink at the finish.

Taken and pecked to pieces, and eaten bit by bit by Silver
Pheasant. The bird wiped his beak several times on the earth,
and for some little time afterwards stood opening and shutting
his beak like a monkey or a human being getting the flavour of
something tasty.

Some of the birds which tried to eat the Timarcha showed no
special signs of finding them unpalatable. It appeared to me
that they finally refused them on account of the hardness of the
exoskeleton. Probably this prevented them getting at the
softer tissues containing the flavour, whether unpleasant or
otherwise.

Larva of Vimarcha tenebricosa.
(A fat bluish-black grub.)

June 15 to 24, 1909. One eaten with apparent relish by
Meerkat, which only delayed seizing it for about two seconds to
rub it in the sawdust and smell it. ‘This was the same Meerkat
that on a previous occasion had eaten Huchelia jacobee and
rejected the Coccinella.

One taken at once by the same Capuchin that had eaten
E. jacobee and rejected Coccinella; but after crushing it between
his teeth and getting the flavour, the monkey at once took it out
in his hands, contemplated it for a few seconds, and moving his
lips the while as if sampling the flavour, then letting it fall,
retired to the back of his cage, salivated and heaved twice as if
going to vomit.

Another Capuchin in the same cage now picked up the crushed
larva, tasted it, and put it down; and neither of the monkeys
touched it again. So I gave it to the Meerkat, which ate it as
greedily as it did the first.

One given to Armadillo was eaten after a good deal of smelling.
A second was eaten without hesitation.

One given to Dent’s Monkey was eagerly taken and tasted,
but almost at once dropped. The monkey did not taste it again,
although he was interested in it and played with it for some
little time.

One given to Mona Monkey, which behaved in much the same
way as Dent’s Monkey, but played with the larva for a longer
time.

One given to Capuchin (sp. a) was taken and chewed up, but

844 MR, R. I. POCOCK ON THE

just as I thought he was going to swallow it, he spat it out with
profuse salivation.

One given to another Capuchin (sp. @) was licked and dropped.

One given to a third Capuchin (sp. 6) was chewed up and
swallowed without any signs of dislike, the larva being not even
taken from his mouth for examination.

Another given to the same monkey was also eaten without any
signs of dislike, although he held it in his hands and licked it
several times before finally putting it into his mouth and chewing
it.

June 24, 1909. Repeated experiments with monkeys.

The two Capuchins (sp. a), the Dent’s and Mona Monkeys
behaved exactly as before. They took the larve, smelt them,
tasted them once or twice, and finally rejected them. The
Capuchin (sp. 6) which had previously eaten two, again ate one
without signs of relish or the opposite. [I then offered a larva
to another Capuchin of the same species (}) and he treated it as
the specimens of the species @ and as the Mona had done, that
is to say smelt it, tasted it, rubbed it in his hands, repeated the
tasting once or twice, and finally dropped it. His behaviour
showed that the difference between the behaviour of the first
example of sp. b, which ate the larvee, and that of the examples
of sp. a, which rejected them, is not attributable to the specific
distinction between the Monkeys as might have been supposed, if
only one specimen of sp. 6 had been available for experiment.

One given to Canadian Jay, taken, pecked, jammed into a
cranny, and repeatedly pecked ; then dropped. When the bird
made no attempt to fetch it, the keeper picked it up and placed
it on the perch, when the bird again seized it, jammed it into a
cranny in the perch, and left it.

One given to Red-backed Shrike was eagerly seized, and after
one or two pecks was left, the bird retiring and wiping his beak
on the bars, as the Canadian Jay had also done.

Two given to Silver Pheasant were taken and pecked, and after
a good deal of rubbing in the earth were eaten.

One given to Prince of Wales’ Pheasant was taken, pecked and
rejected.

One given to Piping Crow was pecked and tasted and rejected,
after a good deal of shaking of the head and wiping of the beak
on the part of the bird. It was then picked up by a Magpie,
which after a taste or two stowed it away under a large stone,
and built up the hole with pebbles.

One given to Buff Laughing Kingfisher was taken and tasted,
but rejected with much bill wiping. Tried and rejected in the
same way by a second specimen of this bird.

One given to Common Laughing Kingfisher was taken and
tasted, but finally rejected.

One given to Dial Bird was finally rejected after a great deal
of pecking and tasting, accompanied by much shaking of the head
and wiping of the bill,

PALATABILITY OF SOME BRITISH INSECTS, 845

One given to White-collared Crow was taken, tasted, carried
about, and finally dropped. This bird refused to take a second
specimen offered immediately afterwards.

One given to Hooded Crow was treated in exactly the same
way as the one above-mentioned was treated by the White-
collared Crow. This Hooded Crow also refused a second
specimen.

One given to Wild Turkey was taken and pecked, but soon
rejected.

SEVEN-SPOTTED LADYBIRD (Coccinella T-punctaia).

July 5, 1909. I offered one to the Capuchin which was the
only one of these Monkeys to eat the 7imarcha-larve, thinking
he might be deficient in tasting powers. He took it at once from
my fingers into his mouth, and crushed it between his teeth;
but, presumably as soon as he got the flavour, removed it frora
his tongue with his fingers, and took no further notice of it.

I offered the remains to a Mona Monkey, but she only Smelt
them and pulled them to pieces, and would not taste them.

July 23 to 31, 1909. One was offered the Capuchin (sp. @)
that had eaten the Huchelia jacobee and Bombus lapidarius on
the previous day, and had so far vefused nothing in the way of
Lepidoptera. He took it from my hands directly, transferred it
to his mouth and crushed it; but instantly took it from his
tongue, wiped it on the perch and left it without a second look.
I then gave the crushed insect to the Meerkat that had eaten
E. jacobee and the Bombus lapidarius. He seized it at once, but
just as promptly spat it out, gave his mouth a wipe with his paw,
and never attempted a second taste.

One given to Vervet Monkey which had just eaten a Timarcha
tenebricosa (see p. 842). She took it, smelt, licked and examined it
thoroughly, rubbed it between her hands, then dropped it to the
floor and took no further notice of it. I had previously offered
this Coccinella to a Chacma Baboon. She smelt it but would not
take it from my fingers.

One given to the Capuchin which on a previous occasion had
tasted and rejected one. He took it, and after a great deal of
smelling, tasting, rubbing between his hands and on the boards
of the cage, finally ate it bit by bit, pulling it into many little
pieces. This Capuchin had just before eaten a Carabus violaceus.

One smelt but refused by three Meerkats. Grabbed by Yellow
Meerkat, tasted, but let go unhurt. Taken by Banded Mongoose,
and eaten after much rubbing in the sawdust, and with many
shakes of the head.

One offered to Grey Lemur, was smelt, taken in the hand and
dropped.

Sept. 20, 1910. One taken and quickly eaten by Meerkat;
but the same animal refused a second specimen.

One taken and rubbed about in the sand and repeatedly bitten,

Proc, Zoou, Soc.—1911, No, LVITL. 58

846 MR. R. I. POCOCK ON THE 2

and ultimately eaten by another Meerkat, but the same animal
refused a second.

One taken in the paws by a Marsh Mongoose, but rejected
after being repeatedly rubbed in the sand and smelt.

One taken by Banded Mongoose and crushed, but rejected with
much head-shaking; swallowed by a second animal also with
much head-shaking.

One refused after being smelt by three Yellow Meerkats.

One taken and licked by Capuchin, but rejected.

One licked but rejected by Red-handed Marmoset.

Another monkey of same species, and a Common Marmoset
refused even to taste it.

July 23, 1909. One examined by Spectacled Thrush, but not
touched. Pecked by Fantailed Flycatcher, which shook his
head and left it. The bird returned three times, however, and
pecked the beetle, but finally gave it up. I then offered it to
a Shama three times in succession, and upon each occasion he
flicked it away and made no attempt to follow it up. Next I
tried the Harmonious Shrike-Thrush. He took it, and after a
good deal of pecking, ate it.

July 31, 1909. Three eaten in succession by cock Silver
Pheasant. The first one he took from my hand, but put it out
of his beak on to the ground. After one or two pecks, however,
he swallowed it. The others he took from my fingers and bolted
entire as if they were grain, exactly as he had previously bolted
the beetles, Pterostichus niger and Ocypus olens.

Sept. 20, 1910. One taken by Pearl-spotted Owl, but dropped
at once.

One taken by a Pekin Robin, which after a few pecks and head
shakes left it and took a drink of water; tasted by another bird
of the same kind, but also left uneaten.

One taken and swallowed, after a deal of pecking about in the
sand and head shaking, by another specimen of Pekin Robin,
which had just previously eaten the grasshopper (Stenobothrus)
and the bug (Therapha hyocyami).

One given to the Dial Bird that had just eaten a Humble Bee
(Bombus agrorum). He took it at once, and after a little delay
swallowed it whole.

N.B. This is the bird that rejected the two White Butterflies
(Pieris brassice and nap?) after tasting them.

One taken but rejected by Masked Wood-Swallow ; then taken
and eaten by Shama.

Two taken and hkolted quickly by the same Shama, which
showed no signs of objecting to the taste, except a single shake
of the head on each occasion after swallowing the beetle.

Although some of these beetles were eaten both by mammals
and birds, there can be no doubt that they were distasteful to the
majority of the animals to which they were offered, even to some
of those that ate them.

PALATABILITY OF SOME BRITISH INSECTS. 847

The interest of the demonstration of the distastefulness of
Coccinella 7-punctata lies in the fact that Coccinellide of various
kinds are numicked in the tropics by insects of other orders, as
well as by spiders.

Order HEMIPTERA.

OxtvE-BROWN Bue (Zropicoris rufipes).

July 21, 1909. One (dead) given to Mona Monkey was eaten
after a great deal of handling, smelling and tasting.

One put on the ground was tackled by Fantailed Flycatcher,
which pecked it some half dozen times. He was then driven off
by a hen Black Tanager, which pecked it and pecked it again, and
then left it. A Syrian Bulbul then flew up and tried it, but after
persevering for some little time gave it up. Then the Tanager
had another attempt, but left it. I then gave the mangled
remains to the Harmonious Shrike-Thrush, and after a little
pecking about he swallowed them.

One (living) eaten with very little delay by Silver Pheasant ;
but put on the ground after being taken from my hand. This
specimen was immature on arrival; 1t moulted in the box, and
was apparently adult when given to the bird.

One (dead) treated in the same way and eaten by the same
bird.

Rep anD Buack Bue (Therapha hyocyami).

Sept. 20,1910. One given to Pekin Robin was at once taken
and ultimately eaten; but the bird took a long time over it,
putting it on the ground after each peck and vigorously shaking
his head before tasting it again. The behaviour of this bird was
exactly the same towards Coccinella 7-punctata (p. 846).

Order DIPTERA.

Bombus-like Fly (Volucella bombylans).

July 26, 1909. One taken by Fantailed Flycatcher but after
being pecked and pulled about for some time, was left. The
Sulphury Tyrant then tried it, but also left it alone after much
pecking. Finally it was taken by Spectacled Thrush, which ate
it after much pecking and wiping in the sand.

One given to Black-headed Sibia was eaten after a great deal
of pecking and breaking up.

These experiments, as Dr. Longstaff reminded me, suggest that
this fly is, at all events to a certain extent, unpalatable. If
future tests should prove it to be so, its likeness to Bombus will
be an instance of Millerian rather than of Batesian Mimicry.

See also below, pp. 854-855,

58*

848 MR. R. I. POCOCK ON THE

Bombus-like Fly (Arctophila mussitans).

See below, pp. 851 and 853.

Fly like a small Bombus (Chilosia illustrata).
See below, pp. 854-855.

Spiny Fry (Lehinomyia ferox).

July 31,1909. One (dead) taken by female Tanager, but after
a good deal of pecking, was left. A Black-headed Sibia then
tried it and finally ate it.

One also eaten by Sulphury Tyrant (see below, p. 855).

Dappy Lone-LeEes (Tipula oleracea).

Oct. 26, 1909. One taken from my hand and eaten readily by
Dial Bird; one taken and eaten, but not so readily, by a second
Dial Bird ; one eaten greedily by Fantailed Flycatcher.

One of these specimens of Tipula was taken twice by the
Harmonious Shrike-Thrush, but was dropped on both occasions.
Another was taken three times by Black-winged Grackle, but was
not eaten.

The rejection of this insect by the Shrike-Thrush, which ate
almost every insect other birds refused, was very surprising.

Fly (Zmpis tessellata).

July 31, 1909. Two (dead). Katen greedily by the Dent’s
Monkey that took the Thanaos tages with avidity (p. 831).

Order HYMENOPTERA.

Tipula-like Ichneumonid (Ophion luteus).
(Nocturnal species, mahogany-red in colour, with
very tough integument.)

Oct. 26, 1909. One taken and tried perseveringly by Fantailed
Flycatcher, but ultimately abandoned. Also tried but soon given
up by Yellow-crowned Hangnest; taken and after a little pulling
about swallowed entire by Dial Bird.

Noy. 7, 1909.—Taken by Black-winged Grackle; but so hard
was the insect that it shot away out of his beak. The bird
pounced on it at once on the sandy floor of the aviary and
ate it; but if the insect had not been very lethargic, or if it
had fallen amongst the undergrowth, it might have escaped
him. Hence probably the significance of its hard slippery
exoskeleton,

PALATABILIVY OF SOME BRITISH INSECTS. 849

Larvee of Saw-fly (Cladius viminalis).

These larve were yellow with black spots. They were sent to
me by Mr. Taylor.

Aug. 19,1910. Refused without tasting by Yellow-crowned
Hangnest, Crested Bulbul, Blue-bird, and Fantailed Flycatcher.

Tasted but rejected by Black-winged Grackle, Harmonious
Shrike-Thrush, Black-chinned Laughing Thrush, and Green
Toucanet.

Taken by Greater Spotted Woodpecker, placed in a hole in a
stump and hammered, but ultimately flicked away and lost.

Two taken and eaten after much pecking and tasting by a
Shama. One eaten fairly readily by a Dial Bird; but another
bird of the same species rejected a specimen after tasting and
flicking it from his beak about twenty times.

Woop-Ant (Yormica rufa).

May, 1910. Taken and eaten with avidity by the following
birds :—Pearl-spotted Owl; Orange-headed Ground-'Thrush ;
Dial Bird; Shama ; Black-headed Sibia ; Blue-bird ; Pekin Robin;
Harmonious Shrike-Thrush ; Spotted Oriole ; Larger Hill Mynah ;
Black-winged Grackle; Yellow-crowned Hangnest; Greater
Spotted Woodpecker.

A Capuchin Monkey also ate one after another, picking them
up in his hands and gobbling them as fast as possible.

Several specimens thrown into a cage containing three Wall
Lizards were tasted by two of them, but rejected at once without
being damaged in any way by the tasting.

Most of the birds showed no signs of objecting to the taste of
the ants, or even of perceiving anything peculiar in their flavour.
The Pearl-spotted Owl, however, shook his head, and the Spotted
Oriole wiped his beak on the perch after eating them. The
Pekin Robin and the Black-winged Grackle wiped the ants upon
their wings, presumably to remove the formic acid. It is inter-
esting to find the same device practised by two species so unlike
one another.

I found that the birds, like the monkey, would eat as many
of these ants as were given to them.

The unavoidable conclusion that these insects are palatable is
rather surprising in view of the frequency with which ants of
different kinds are mimicked in the tropics by Orthoptera, Coleo-
ptera, and other insects, as well as by spiders. Nevertheless, it
corroborates the opinion put forward by McCook and amplified
and endorsed by myself in 1909 *, before these experiments were
made, that ant-mimicry is mainly serviceable as a protection
against the predatory Hymenoptera of the family Pompilide, which
provision their nests with Arthropoda of various kinds, excepting
ants, and are certainly the direst enemies that spiders possess.

* Journ. Linn. Soc., Zool. xxx. pp. 265-268.

850 MR. R. I. POCOCK ON THE

Saw-rry (Allantus arenatus).

July 21,1909. One eaten by Mona Monkey fairly readily ; by
nee readily ; by the Capuchin which on the previous day
had refused the Malacoderm Beetle (Rhagonyche fulva); smelt,
but not tasted by Lion Marmoset.

One eaten fairly readily by Harmonious Shrike-Thrush ; by
Shama readily; by Silver Pheasant; refused without tasting by
Wild Turkey.

Honey-BEE (Apis mellifica). (Workers.)

May 8, 1911. One offered to Silver Pheasant was taken from
the forceps but immediately flicked away; the bird persevered,
however, and after much pecking and flicking ¢ about of the insect,
and wiping his bill on the ground, finally ate it.

One offered to Bornean Fire- backed Pheasant was inspected
carefully but rejected untasted.

One given to Pekin Robin was taken at once, but was quickly
flicked away. When pursued, however, by other birds in the
cage, the Pekin Robin pounced on the bee again and flew away
with it. Whenever he got a moment’s peace, he put it on the
eround, pecked and flicked it about, wiping it now and again in
the sand and repeatedly shaking his head. At length he flew to
a branch, and holding the bee against it with his foot, pulled it
in two pieces, dropping one piece to the ground. He still
persevered with the other piece, however, but an finally lost sight
of him and do not know whether he ate it or not.

One given to a Cayenne Tanager was taken and chewed for a
long time ; ; the remains, however, were finally jammed into a
banana and left.

One taken by a Blue Tanager which, however, allowed himself
to be robbed without resistance or flight by a Maroon Tanager.
This bird, after a deal of mastication, ate the bee.

One given to Wall Lizard was eagerly seized, but was left after
one or two attempts.

Another was twice darted at by another lizard of this species,
but was left alone the moment the lizard touched it. It was then
boldly seized by a third lizard, which with one bite disabled the
bee by crushing the head and thorax. This lizard persevered for
about seven minutes, biting at the bee, but stopping after each
bite to lick his mouth with his tongue and Pub it against the moss.
Finally he gave it up and went away.

Two Bluebottles(Calliphor a vomitoria) “ a Hover-fly (Syrphus)
given as a check experiment were seized and eaten in a few
seconds by the same lizards.

Humpte Bes (Bombus agrorum).
(See also infra, p. 853.)

Oct. 26, 1909. One eaten with avidity by Capuchin and by
Meerkat.

PALATABILITY OF SOME BRITISH INSECTS. 851

One given to Collared Jay-Thrush, which pecked it about and
scraped it in the sand for along time, wiping his beak in the
intervals, and ultimately left it. It was then picked up by a
White-crested Jay-Thrush, which treated it for some time in the
same way, but at last ate the mangled remains. This same bird
then took a specimen of the mimetic fly Aretophila mussitans, but
made just the same fuss over the eating of it as he had in the
case of the bee.

Sept. 18,1910. One taken at once by Dial Bird, and after a
good deal of pulling about, pecking and wiping in the sand, was
eaten. This bird had just previously eaten a small Tortoiseshell
Butterfly, and he took about the same time to finish off the one
insect as the other.

Sept. 20,1910. One offered to Dial Bird was taken at once and
eaten with very little delay, after being wiped once or twice in
the sand. The bird flew away with a second specimen and I did
not see what became of it; but he returned to me, and I had
difficulty in keeping him away from the bees with which I was
experimenting with other birds.

This Dial Bird was the one that ate the same species of Humble
Bee two days previously.

Humsie Bee (Lombus ? joncellus).

July 31, 1909. One offered alive to Mona Monkey was
snatched at once and eaten bit by bit.

Houms1e BEE (Bombus ? terrestris).

July 31,1909. One (dead) taken by Brazilian Hangnest and
pecked to pieces, the bird holding it the while in his foot against
the perch. The pieces pecked off were dropped about the cage
and not eaten.

HuMBLe BEE (Lombus lapidarius).

May 31, 1909. One dead specimen given to the Meerkat was
eaten bit by bit, after being rubbed im the sawdust by the
animal’s paws.

One dead specimen given to Capuchin (Cebus sp. a) was taken
in the hands and eaten bit by bit, just as the Monkey would eat
a piece of hard biscuit or sugar. Neither of these mammals
showed any signs of disliking the taste of the bees; quite the
contrary. Their molar teeth are evidently much better adapted
for crushing the chitinous exoskeletons than are the beaks of the
birds that tasted them.

One dead specimen offered to Syrian Bulbul was taken after
about a minute’s inspection. The bird pecked it and pulled it
about for at least five minutes and dodged away with it from
other birds that chased him. He grew, however, less and less

852 MR. R. I. POCOCK ON THE

keen, and ultimately allowed a female Black Tanager to rob him
of it. The Tanager behaved in just the same way, pecking and
pulling it about and breaking it to pieces, but gradually losing
her interest. At last she picked up a piece of the thorax and
flew to a bush with it, leaving the remainder on the ground. I
could not see what became of the piece she flew away with, but
she emerged from the bush without it, and wiped her beak on
a perch. She made no attempt to go back to the bits on the
ground. A Sibia tried these, but after a peck or two left them,
and no other insectivorous bird in that compartment took the
least notice of them. So I picked up the abdomen and gave it to
the Harmonious Shrike-Thrush which had just finished off the
example of B. hortorwm, mentioned below, and he ultimately ate
it after a great deal of pecking and pulling about.

Houmste Bre (Bombus hortorum).

May 31,1909. One living example fell to the ground of the
aviary when first liberated. Two Fantailed Flycatchers flew
down to it at once, but although interested would not touch it ;
while they were hesitating the bee took wing and escaped, none
of the birds in the aviary making any attempt at pursuit.

One dead specimen offered to a Shama. She allowed me to
hold it close to her beak, but would not touch it. None of the
other birds in the aviary would notice it when thrown to the
ground, though on a previous occasion they had shown great
eagerness in seizing dead butterflies. I then gave it through the
bars to the Harmonious Shrike-Thrush in the next compartment.
After pecking and pulling it about for six or seven minutes, he
ultimately ate it.

July 31, 1909. One sniffed at but rejected by two Meerkats ;
taken by a third and eaten.

One pecked and flicked away by Black-headed Sibia, by Shama.
and also by Sun-Bittern, each making two or three attempts.
Then carried off by female Black Tanager, but dropped to the
floor, where a North American Cat-bird tried it once or twice, but
gave it up. (The remains were now too mangled to be useful for
further experiment.)

T could not induce the Fantailed Flycatcher to take any notice
of this bee.

Conclusion. These experiments indicate that the Humble Bees
used for the tests were much more palatable to the mammals than
to the birds. With the exception of the one example of b. lapi-
darius which was smelt and left untouched by two Meerkats, all
the bees offered to the Monkeys and Meerkats were eaten without
any kind of dislike of the flavour being evinced. The Meerkat
that rubbed the B. lapidarius in the sawdust did so, I suspect, to
remove some substance offensive to his sense of smell. On the
other hand, of the birds to which the bees were oftered only three

PALATABILITY OF SOME BRITISH INSECTS. 853

ate them, namely a Dial Bird, a Jay-Thrush, and a Shrike-Thrush.
The Dial Bird ate one quickly with only one or two wipes in the
sand. In the other cases there was a great deal of pecking and
wiping before the insects were finally disposed of. From the
behaviour of the birds there could be no doubt that there was
something in the bees not to their liking, even to those that
ultimately ate them. The Bulbul, Sibia, and Tanager were
obviously keen to eat them, and gave them the fullest possible
trial before finally rejecting them ; but whether it was the hairs,
or the hard chitin, or the favour, or a combination of them that
made the insects unpalatable, I do not know.

Further experiments demonstrating the distastefulness of
Humbie Bees to birds of different kinds are given in the following
section :—

Experiments to test the significance of the resemblance between
Humble Bees (Bombus) and the Flies Arctophila nussitans,
Volucella bombylans, and Chilosia illustrata.

Bombus agrorum and Arctophila mussitans.

Oct. 26, 1909. Offered Bee to a Lion Marmoset which was
busily catching house-flies and bluebottles in his cage. He
looked at it, but would not touch it. I then offered the fly, but
he also refused to touch it. He did not, however, hesitate to take
a Red Admiral offered a moment afterwards.

Offered Bee to Leach’s Laughing Kingfisher. He took it at
once, but soon flicked it away. Six times in succession he took it
from my fingers and dropped it on each occasion. | could not
induce him to take it again. Instead he started pecking my
fingers. Thereupon I offered him the fly, and he just as reso-
lutely refused to take it.

Offered Bee to Kagu, a New Caledonian Rail. He inspected
it, and after a little hesitation tasted it. But he would not touch
it again; and when offered the fly, refused that likewise.

Offered Bee to Central American White-browed Partridge.
He took it without hesitation, but after a peck or two left it and
went away. I then threw it to him, and he tasted it again ; but
would not eat it. I then threw him the fly, but he would not
touch it.

A Douracouli (a South-American monkey) ; a Honduras Turkey ;
a cock and a hen Reeves’s Pheasant, and three hen Silver
Pheasants refused to touch both bee and fly, though they
inspected them intently for a few seconds.

Sept. 20, 1910. Bee offered to Hoopoe was taken at once and
tasted without being crushed, but was then left on the ground
uneaten. The bird refused the next one I offered, and then
refused to take the fly, although he stretched his head towards
it and inspected it.

Bee offered to Yellow-crowned Hangnest, which took it at once,
but soon dropped it. A second time he took it, and dropped it.

854 MR, R. I. POCOCK ON THE

The third time it was offered he refused it, and immediately after-
wards refused the fly.

Bee offered to Sulphury Tyrant. I importuned the bird into
taking it from my fingers no fewer than eight times, and each
time he flicked it away. The ninth time he refused to take it,
and then refused the fly.

Bee offered to Black-winged Grackle, which took it at once, but
dropped it. Twice more he took it and the last time flew a short
distance away and persevered with it for about three-quarters of
a minute, then leaving it returned to me; he refused the next
bee I offered, and then refused the fly.

Bee offered to Silver Pheasant was at once taken, put on the
ground, pecked and crushed almost past recognition, but left
uneaten. The bird then took from my fingers three more speci-
mens in succession, but dropped them uncrushed from his beak
at once. The fifth he looked at, but would not touch, and then
also refused the fly after inspecting it.

The experiments described above with the Lion Marmoset, the
Douracouli, the Turkey, and the Reeves’s and Silver Pheasants,
which would not touch either the bee or the fly after some seconds
of intent inspection, do not prove that the bee was known to be
distasteful, and that the fly was rejected in consequence. That
may be the explanation, The Douracouli, however, is nocturnal
and probably does not naturally feed upon diurnal-flying insects.
In the case of the Marmoset, the experiment does, however,
suggest very forcibly that the Arctophila was not recognised as
closely allied to the bluebottles the animal was hunting. The
other experiments speak for themselves.

Bombus hortorum, Volucella bombylans, and
Chilosia illustrata.

July 31,1909. Offered living Bombus hortorum to the Bra-
ailian Hangnest that had just pulled the dead Lombus terrestris
to pieces. He took it directly, but instantly flicked it away and
wiped his beak. The bee then crawled up the bars of the cage,
and he again pecked and flicked it away. It was now too injured
to crawl although still alive, so I picked it up and offered it in
my fingers. He took it again and flicked it away. Twice more
the trial was made, with the same result, although he was
patently tiring of the trials. The next time he refused to
touch it after inspection. I then substituted a dead Volucella
bombylans. We inspected it, but did not touch it, and hopped up
to the top perch.

I then offered the nearly dead Bombus to another specimen of
the same bird. He took it from my fingers three times in
succession, and each time flicked it away. The fourth time he
refused to touch it. I then substituted the same specimen of
Volucella bombylans, but after looking at it he would not take it.

PALATABILITY OF SOME BRITISH INSECTS. 855

I then again offered the Bombus to the first Hangnest. He
took it and flicked it away, and immediately afterwards refused
to touch the Volucella.

Next day I offered the first Hangnest a Bombusagain ; he took
it from me three times, and flicked it away without attempting
to eat it, but immediately afterwards took Chilosia illustrata and
ate it.

The second Hangnest took a dead Lombus, and flicked ite away,
and then ate Chilosia ilustrata, but refused immediately after-
wards to touch a live Bombus hortorwm.

Offered Bombus hortorum to North American Cat-bird, which
came up to me on seeing other birds being fed. He pecked it
several times, but flicked it away and gave it up. During the
next quarter of an hour I could not induce him to touch either
Volucella bombylans or Chilosia illustrata.

T then offered the Bombus on the forceps to a Sulphury Tyrant.
He pecked and flicked it away several times, then left it, and
refused it when offered again. I then offered him the Volucella
bombylans both in the forceps and by throwing it to him on the
ground, but he would not touch it. After a little hesitation,
however, he took an Hchinomyia ferow from the forceps and ate
it, and then took and ate Chilosia illustrata. I then offered him
Bombus hortorum again, and he took it but soon rejected it, and
immediately afterwards refused to touch Volwcella bombylans.

One Bombus hortorwm ottered to a Shama, which pecked it once
or twice, and flicked it away each time. He then refused to touch
the specimen of Volucella bombylans.

Tried the experiment with another Shama, which behaved in
exactly the same way towards the bee, and would not afterwards
touch the Volucella bombylans.

One Bombus hortorwm offered to Silver Pheasant was taken at
once, but left after some pecking and tasting. Then without
hesitation he took Chilosia wlustrata from the forceps and ate it ;
and promptly tried the Bombus again as it lay on the ground,
but would not eat it. Immediately afterwards he eagerly ate an
Ocypus olens and three specimens of Péerostichus (see pp. 837-838).

I made one Volucella bombylans do duty for all the experiments
described above and had it intact at the end. It was not pecked
by any of the birds, presumably because I never offered it to one
until he had tried Bombus hortorwm a sufficient number of times
to reject it as unpalatable ; and there is no doubt in my opinion
that they did not distinguish between the bee and the fly.
Although Chilosia illustrata is also very like Bombus, the difference
in size is well marked. I suspect that in this circumstance lies
the explanation of the birds not confusing this species of fly with
the bee. They could judge the difference in size quite easily,
because the insects were held at the same distance from them.

856 MR. R. I. POCOCK ON THE

List of THE MAmmats, Brrps, AND REPTILES USED FOR THE
EXPERIMENTS.

MAMMALS.

Mona Monkey (Cercopithecus mona), Nigeria. Diana Monkey
(Cercopithecus roloway), Gold Coast and Guinea. Dent’s
Monkey (Cercopithecus denti), Ituri Forest. Vervet Monkey
(Cercopithecus pygerythrus), Cape Colony. Yellow Baboon
(Papio sphinx), Nigeria. Ceylonese Macaque (J/acacus
puleatus), Ceylon.

Although feeding mostly upon fruits, roots, and vegetables of
various kinds, all the Monkeys of the Old World eat insects as
well.

Capuchins (Cebus, spp. 2).

Several immature specimens, belonging to undetermined species

inhabiting the forests of the northern parts of South America.

Douracouli (Wyctipithecus trivirgatus).
A nocturnal Monkey from the Amazons.

Lion Marmoset (Zeontocebus rosalia). Pinché Marmoset
(Leontocebus adipus). Red-handed Marmoset (Leontocebus
rufimanus). Common Marmoset (Callithrix jacchus).

Although vegetable feeders in the main, the South American
Monkeys and Marmosets seem more addicted to an insect diet
than the Monkeys of the Old World.

Grey Lemur (Hapalemur griseus). Crowned Lemur (Lemur
coronatus). Black Lemur (Lemur macaco). White-fronted
Lemur (Lemur fulvus albifrons). Mongoose Lemur (Lemur
mongoz).

Lemurs inhabit Madagascar. They do not appear to be partial
to insects.
Suricate or Meerkat (Suricata swricatta).

Cape Colony. Feeds on small animals of various kinds and
particularly insects and their grubs (W. LZ. Sclater).

Yellow Meerkat (Cynictis penicillata).
Cape Colony. Feeds on small birds, mammals, eggs, and insects

(W. L. Sclater).

Banded Mongoose (Crossarchus fasciatus).
South and East Africa. Feeds on insects, fruits, seeds, eggs,
snails, etc., according to Bohm.

Common Indian Mongoose (J/wngos mungo), from India, and
McCarthy’s Mongoose (Ifungos fulvescens), from Ceylon, live on
small mammals, birds, reptiles, insects, and fruit. The White-
tailed Mongoose (Mungos albicauda), from Africa south of the

PALATABILITY OF SOME BRITISH INSECTS. 857

Sahara, does not, so far as is known, differ in diet from the other
species just mentioned.
Marsh Mongoose (Mungos galera).
West and South Africa. An amphibious species feeding
mainly it is alleged upon crabs, fishes, frogs, and insects.
Grison (Grison furax=Galictis vittata).
A musteline carnivore from the Argentine, feeding upon
small mammals and birds but also fond of fruit.
Common Armadillo (Dasypus villosus).
Argentine. Feeds on insects, grubs, worms, carrion, and
vegetable matter.

BIRDS.
Cape Robin-chat (Cossypha caffra).
Range. East Africa to Cape Colony.
Food. Chiefly insects, spiders, and worms; also berries and
small fruit (Sclater d: Stark).
Common Thrush (Vurdus musicus).
Range. Palearctic Region, locally migratory.
Food. Insects, worms, fruit, etc.
Orange-headed Ground-Thrush (Geocichla citrina).
Range. The Himalayas up to 5000-6000 ft., Assam and
Tenasserim.
Blue Rock-Thrush (Geocichla (Monticola) cyanus).
Range. From South Europe and North Africa to Turkestan,
Tibet, the Himalayas, and Burma.
Common Rock-Thrush (Geocschla (Monticola) saxatilis).
Range. C. & S. Europe to C. Asia, N.E. Siberia and N. China.

Wood-Thrush (Hylocichla mustelina).
Range. Eastern North America, Central America to Guatemala.
Dial Bird (Copsychus saularis).
Range. Ceylon, India, ascending the Himalayas up to 5000 ft. ;
Burma and Tenasserim.
Shama (Crttocincla macrura).
Range. Ceylon, India and Burma.
Blue-bird (Szalia sialis).

Range. Eastern North America to a little west of the Missouri
River.

Food. Insects of various kinds ; also ripe fruits.

American Cat-bird (Galeoscoptes carolinensis).

Range. South-eastern United States to the Missouri, migrating
southwards in the winter.
Food, Insects, fruit and seeds,

858 MR. R. I. POCOCK ON THE

Mocking Bird (Wimus polyglottus).
Range. Southern United States from the Atlantic to the high
central plains ; locally migratory.
Food. Insects and fruit.

Cuban Mocking Bird (Mimus orpheus).
Range. Jamaica, Porto Rico, Haiti, Cuba.

Saturnine Mocking Bird (imus saturninus).
Range. Brazil.

Great Tit (Parus major).
Range. Widely distributed in the Palearctic Region. Locally
migrating but mostly resident.
Food. Insects and seeds.

Pekin Robin (Liothrix luteus).
Range. Himalayas from Simla to Bhutan ; extending also into
China ; resident.
According to E. W. Oates the food of this bird consists of
berries, fruit, seeds, and insects.

Pied Grallina (Grallina australis).

Range. Australia, generally distributed.
, Food. Insects (Gould).

White-eared Bulbul (Pycnonotus lewconotus).

Range. Persia; Sind, the Punjab, the N.W. Provinces of
India, and Central India as far east as Hoshargabad.

White-cheeked Bulbul (Pycnonotus leucogenys).

Range. Afghanistan ; the Himalayas from Murree to Bhutan,
up to 7000 ft.

Red-vented Bulbul (Pycnonotus hemorrhous).
Range. Ceylon; India roughly to the foot of the Himalayas.
According to E. W. Oates the Indian species of Bulbuls feed
chiefly upon fruit.

Syrian Bulbul (Pycnonotus xanthopygus).
Range. N.E. Africa, Arabia, Palestine, Cyprus.

Black-crested Bulbul (Otocompsa flaviventris).
Range. Nepal to Cochin China.

In the course of my experiments I noticed that Bulbuls of
different species were very keen on butterflies ; of beetles and
crawling insects generally they took little if any notice; but the
moment a butterfly was let loose in the aviary they were all on
the move. From this I infer that they are great butterfly-
hunters in their own countries,

PALATABILITY OF SOME BRITISH INSECTS. 859

Orange-headed Laughing Thrush (7rochalopteron
erythrocephalumy).
Range. Himalayas, from Chamba to Nepal up to 7000 ft.

Black-chinned Laughing Thrush (Zvochalopteron nigrimentum).

Range. Himalayas from Nepal to Assam (7000 ft.).

According to E. W. Oates the food of the Laughing Thrushes
(Trochalopteron) is the same as that of the Jay-Thrushes
(Garrulaz).

Spectacled or Melodious Jay-Tbrush (Trochalopteron canorwm).

Range. China; Shanghai, Amoy, Fokien, Chekiang.

Black-headed Sibia (Sibia capistrata).
Range. Himalayas from Hazira to Bhutan, 5000-8000 ft. ;
resident.
Collared Jay-Thrush (Garrulaz picticollis).
Range. China: Chekiang, Fokien.

White-crested J ay-Thrush (Garrulax leucolophus).

Range. Himalayas to Assam and Burma in the hill-tracts.

According to E. W. Oates the Indian species of Garrulax feed
upon every sort of insect and smaller reptiles, and probably also
on fruit.

Grey Struthidea (Struthidea cinerea).
Range. South-eastern Australia; resident.
Food, Insects, particularly beetles.

Spotted Oriole (Oriolus maculatus).
Range. Sumatra, Java, Borneo.

Harmonious Shrike-Thrush (Collyriocinela harmonica).

Range. Australia; N.S. Wales and 8. Australia.
Food. Insects (Gould).

White-eyebrowed Wood-Swallow (Artamus superciliosus).

Range. Interior of South Australia.
Food. Insects (Gould).

Masked Wood-Swallow (Artamus personatus).

Range. South Australia, locally migratory.
Food. Insects (Gould).

Red-backed Shrike (Lanius collurio).

Range. Europe, migrating in the autumn and winter into
Western India and to South Africa.
Food. Insects; small birds etc.

Fantailed Flycatcher (Lhipidura tricolor).

Range. Australia, widely distributed.
Food. Insects of various kinds (Gould).

860 MR. R. I. POCOCK ON THE

Garrulous Honey-eater (yzantha garrula).

Range. South Australia, Tasmania,
Food. Honey and insects (Gould).

Black Tanager (Z’achyphonus melaleucus).

Range. Costa Rica through Panama, Venezuela, Ecuador to
Bahia.

Scarlet Tanager (Rhamphocelus brasilius).
Range. South-eastern Brazil.

Cayenne Tanager (Calliste cayana).
Range. Guiana, Venezuela, Ecuador, Peru.

Green Hangnest (Ostinops viridis).
Range. Guiana, Brazil, Ecuador.

Yellow Hangnest (Cassicus persicus).
Range. Trinidad, Guiana, Ecuador, Bolivia, Brazil.

Common Hangnest (/cterws vulgaris).
Range. Colombia, Venezuela.

Brazilian Hangnest (Jcterws jamaicat).
Range. North Brazil.

,Yellow-crowned Hangnest (Jeterus chrysocephalus).
Range. Guiana, Venezuela, Kcuador, Brazil.

Larger Hill Mynah (Gracula intermedia).

Range. India: the south-eastern Central Provinces, the lower
ranges of the Himalayas from Kumaon to Assam, thence into the
Malay Peninsula.

Small Hill Mynah (Gracula religiosa).

Range. Ceylon and Southern India.

According to EK. W. Oates these two species of Mynah are
resident or only locally migratory and live exclusively upon
fruit. ,

Chinese Mynah (Acridotheres cristatellus).

Range, China: Shanghai, Hainan, Formosa; Philippine
Islands.

Pied Mynah (Stwrnopastor contra).
Range. Central and South India to Assam and Burma.

Javan Pied Mynah (Sturnopastor jalle).
Range. Sumatra, Java, Borneo.

Black-winged Grackle (Giraculipica melanoptera).
Range. Java.

PALA'TABILITY OF SOME BRITISH INSECTS. 861

Spotted Bower Bird (CAlamydodera maculata).
Range. New South Wales.
Food. Principally fruit and grain (Gould).
King Bird of Paradise (Cictnnurus regius).
Range. New Guinea.
Food. Fruit and insects.
Canadian Jay (Perisorews canadensis).
Range. Canada and the Northern States of the Union.
Food. Insects ; eggs, flesh ; leaves of fir trees (Audubon).
Hooded Crow (Corvus cornix).
Range. Palearctic Region.
Food. Omnivorous (eggs, carrion, young birds, etc.).
White-collared or Pied Crow (Corvus scapulatus).
Range. Atvica south of the Sahara.
Food. Omnivorous, with partiality for flesh food.
White-backed Piping Crow (Gymmorhina leuconota).
Range. 8S. Australia, New South Wales.
Food. Mostly insects (Gould).
Long-billed Butcher Crow (Oracticws destructor).
Range. Australia.
Food. Chiefly insects.
Sulphury Tyrant (Pitangus sulphuratus).

Range. Guiana, Ecuador, Peru, Brazil.
Food. Mostly insects and animal food of various kinds as well

as fruit.
Greater Spotted Woodpecker (Dendrocopus major).
Range. Palearctic Region.
food. Insects.
Common Laughing Kingfisher (Dacelo gigantea).
Range. New South Wales and South Australia.

Leach’s Laughing Kingfisher (Dacelo leachit).
Range. North-east coast of Australia.

Buff Laughing Kingfisher (Dacelo cervina).
Range. Kast and North Austvralia.
Food. These great Kingfishers feed mainly upon reptiles and
insects, but also upon rats and inice.

Hlate Hornbill (Ceratogymna elata) and
Black Hornbill (C. atrata).

Range. W. Africa, Nigeria, etc.
Food. Insects ; snakes, small mammals, ete.

Proc. Zoou. Soc.—1911, No. LIX. 59

862 : MR. R. I. POCOCK ON THE

Ground Hornbill (Bucorax abyssivicus).

Range. North Africa south of the Sahara. ny
Food. Insects, snakes, frogs, lizards (Stark and Sclater writing
of the closely allied southern species B. caffer).

Hoopoe (Upupa epops).

Range. Southern Palexaretic Region from Scandinavia and the
British Islands to Japan, migrating in winter to North Africa,
Arabia and India.

Food. Ground insects, beetles, grasshoppers and ants.

Cartagenian Motmot (Momotus subrufescens).
Range. From Panama, Colombia, and Venezuela to Matto Grosso.

Great Barbet (egalema virens).
Range. China and Upper Burma.

Levaillant’s Barbet (Zrachyphonus caffer).
Range. 8. Africa, Natal, the Transvaal, Rhodesia, ete.
Food. Fruits, berries, leaves, and insects such as termites

(Stark & Sclater).
Green Toucanet (Aulacorhamphus sulcatus).
Range. Venezuela; Colombia.

Pearl-spotted Owl (Glaucidiwm perlatum).
Range. Africa south of the Sahara.
Food. Mostly insects (grasshoppers, termites); also mice and
lizards (W. L. Sclater).
White-eared Scops Owl (Scops leucotis).
Range. Africa south of the Sahara to the Orange River.
Food. Chiefly insects, like grasshoppers; also rats and mice

(W. L. Sclater).
Prince of Wales Pheasant (Phasianus principalis).
Range. North-western Afghanistan and North-east Persia.

Reeves’s Pheasant (Phasianus reevesit).
Range. Mountains of Northern and Western China, extending
as far east as Kiu-Kiang.
Elliot's Pheasant (Calophasis elliott).
Range. Mountains of South-eastern China.

Silver Pheasant (Genneus nycthemerus).
Range. South China, Fokien and Chekiang.

Vulturine Guinea Fowl (Aerylliwm vulturinum).

Range. Kast Africa from the Pangani River westwards to
Kilimanjaro and northwards to Somaliland.

PALATABILITY OF SOME BRITISH INSECTS. 863

Pucheran’s Guinea Fowl (Guttera pucherani).
Range. Kast Africa: Zanzibar to the Tana River and thence
westwards into the interior.
N. American Wild Turkey (Meleagris americana).

Range. Formerly widely distributed in the United States of
America. Not migratory.

food. Beechnuts, acorns, berries, green-shoots, etc. ; also grass-
hoppers, and other insects (Bendire).

Honduras Turkey (Jeleagris ocellata).

Range. Central America: Guatemala, Yucatan, Honduras.
Food. Probably of a similar nature to that of JZ. americana.

Long-tailed Partridge (Dendrortyx leucophrys).
Range. Highlands of Guatemala and Costa Rica (Ogilvie-
Grant).
Brush Turkey (Catheturus lathami).
Range. North-east and Kast Australia.

Crested Curassow (C'raa alector).
Range. Northern part of South America: British Guiana,
Colombia, Rio Negro, ete.
Globose Curassow (Crax globicera).
Range. Central America: Western Mexico to Honduras and
Cozumel Island.
Yarrell’s Curassow (Craw carunculata).
Range. South-eastern Brazil from Rio Janeiro to Bahia.

Red-tailed Guan (Ortalis ruficauda).
Range. Venezuela and the island of Tobago.
Most Game-birds, especially when young, eat insects as well as
grain, nuts, and green-food.

Australian Bustard (Hupodotis australis).

Range. South and Western Australia.
Food. Seeds, vegetables, grasses, and insects (G'owld).

Vigors’s Bustard (Otis vigorsit).
Range. 8. Africa: Cape Colony, Natal, ete.
Food. Seeds, insects, small reptiles (Stark dg Sclater).

Ludwig’s Bustard (Otis ludwig?).

Range. 8. Africa: Cape Colony, Natal, Orange River Colony,
S. Transvaal; partially migratory within this area.

Food. Mostly beetles, caterpillars, and other insects (Stark &
Sclater).

The food of Bustards is probably much the same everywhere.
The diet is essentially mixed, and consists of grain, green-shoots and
leaves insects, smal] mammals (mice) and reptiles,

864 PROF. E. B, POULTON ON THE

Two-striped Thickknee (Hdienemus bistriatus).
Range. Mexico through Central America to Venezuela
Food. Insects, worms, snails, ete.

Trumpeter (Psophia crepitans).
Range. Brazil.
Food. Fruits, seeds, insects.

Cariama or Seriema (Cariama cristata).
Range. South-east Brazil.
Food. Reptiles and smal! mammals for the most part.

Abbott’s Rail (Hallus abbotti).
Range. Assumption Island.

Black-tailed Water-hen (Zribonyx ventralis).
Range. Australia, south of the 25th parallel ; locally migratory.

Kagu (Rhinochetus jubatus).
Range. New Caledonia.

Sun-Bittern (Hurypyga helias).
Range. Northern countries of the Neotropical Region.
Food. Mostly insects.

REPTILIA

The Green Lizard (Lacerta viridis), from Central and Southern
Europe; the Wall Lizard (Lacerta muralis), from Central and
Southern Europe; the Filfola Wall Lizard (LZ. muralis filfolensis),
from Filfola, near Malta; Dugés’s Lizard (Lacerta dugesii), from
Madeira; the Sand Lizard (Lacerta agilis), from North and
Central Europe ; and the Black-spotted Lizard (Algiroides mgro-
punctatus), from Dalmatia, feed mainly upon insects, worms, and
small slugs.

Glass Snake (Ophisaurus apus).

South-eastern Europe. Feeds on small mammals, reptiles,

slugs, ete.

Notes upon some of the above described Huperiments by
Prof. E. B. Poutton, F.R.S., F.Z.8.

Pages 815-820.

The experiments on the Pierinw support the conclusion that
the perfection of the under surface procryptic resemblance affords
a true criterion of the degree of palatability.

P. brassice, with its conspicuous gregarious larva, and imago
larger and less cryptically coloured than the other three species

PALATABILITY OF SOME BRITISH INSECTS. 865

(although nearly the same as P. rape in this respect), was
distinctly the least palatable of the four. On the other hand,
P. napi and EF. cardamines, in which the cryptic resemblance is
carried to its highest pitch, appeared to be most palatable; but
a larger number of experiments is greatly to be desired.

The results obtained in the two former species are of much
interest in relation to the experiments upon J/elanargia galathea

(p. 827).

Pages 820-822.

The evidence that J’. wrticee is not very palatable agrees with
my own experiments * with a Marmoset; and I obtained the
same results with V. 7o when offered in considerable numbers to
lizards. It is probable that the procryptic under surface of the
Vanessas is chiefly related to the attacks of mammals and of very
hungry birds during the long hibernating period. The special
interest in the eye-spots of V. io manifested by the Syrian Bulbul,
accords with previous observations on other insects and other
insect-eaters, Reptilian as well as Avian ft.

Pages 823-825.

The evidence of a certain amount of unpalatability in
Brenthis (Argynnide) is consistent with the degree of proeryptic
defence attained in this genus. It is also of much interest in
connexion with the experiments on Avaschnia levana, the early
or levana form of which is probably a mimic of the species of
Brenthis. The examples of the Araschnia tested by the author
(pp. 823-824) were of the form prorsa, belonging to the later
brood, and generally looked upon as mimics of the White Admiral
(Limenitis sibylla), which appears upon the wing at about the
same period. The experiments here recorded prove that the
mimic is certainly unpalatable to several birds, and support the
conclusion that the resemblance is Miillerian or Synaposematic.
The evidence, so far as it goes, points indeed to the inference
that Araschnia is more unpalatable than its Brenthis model. A
few experiments on the imago of L. sibylla made by Mr. Pocock
in the summer of 1910, also indicated that the prorsa form is
more unpalatable than the Limenitis. There is, however, nothing
improbable in a Miillerian mimic being more highly protected than
its model. The role of model is related to many characteristics,
and relative abundance, conspicuousness, and extent of range
may play their part as well as relative unpalatability. Thus it is
probable, from its habits and flight, that the Eastern Kuropean
Neptis lucilla, W.V.(=sappho Pall.) is more distasteful than its
Limenitis models, but the latter are widespread and abundant
species, and it is reasonable to suppose that the memories of
European insect-eating animals are more deeply impressed by
their pattern than by that of the Veptis.

* Trans. Ent. Soc. Lond. 1902, p. 442.
+ ‘Essays on Evolution ’ (Poulton), 1908, p. 210: see also p. 326,

866 PROF. E. B. POULTON ON THE

Pages 825-827.

The desirability of experiments upon the palatability of the
genus Melitea was suggested by the study in 1908 of a collection of
butter flies from the Tian Shan or Celestial Mountains in Western
Mongolia, Small as it was, the collection was sufficient to show
that Melitea is a dominant element in the insect fauna of the
locality. The large “ Skipper,” Hesperia antonia Spey, was also
abundantly represented, and I was at once struck with the marked
resemblance which its under surface would bear in the position of
rest to that of the species of Melitea. The striking feature of the
latter genus is supplied by two black-bordered orange bands
which cross the hind wings and stand out conspicuously against
the cream ground-colour. These two bands, the outer with its
festooned, the inner with its irregular borders, present a highly
characteristic appearance. ‘The small portion of the fore wing
under surface exposed in the position of rest conforms, as is
usual in butterflies, to the pattern of the hind and appears as a
slight extension of its area. In spite of differences in detail, the
two orange bands of the Hesperid closely resemble those of
Melitea, and in all essential respects the exposed under surface of
the former reproduces that of the latter. In the Skipper the
outer margin of the outer band is cut mto internervular con-
cavities, while the inner bulges into corresponding convexities:
in the Nymphaline, concavities are seen along both borders. The
orange of the bands and the tint of the ground-colour—white
between the bands, greyish elsewhere—are also much paler in the
Skipper, but the orange pigment is probably quite different from
that of Melitea and may rapidly fade. It is also interesting to
note that the orange bands of the under surface are represented
by black bands on the upper surface of the Skipper but by orange
bands on that of the Nymphaline. The allied Hesperia side Esp. Ls
with golden bands, is doubtless a co-mimie with H. antonia, while
im 2 fbr closely related species, 7. carthami Hiibn., the dark
bands have gained a bronzy greenish or yellowish tinge, probably
indicating the kind of variation out of which the pattern of the
two first-named species was produced by selection.

Probable evidence that Melitwa is a specially protected genus is
supplied by the well-known habits of the three British species
aurinia (=artemis) Rott., cinwia L., and athalia Rott. All are
known to be gregarious in the larval state, and so abundant in
confined localities that they may be described as gregarious in
the perfect state also. All are slow-flying and conspicuous on the
wing and at rest between the flights, while individuals have been
observed to “sham death” when captured. Putting all the facts
together, it appeared probable that we have an interesting addition
to the list of mimics among the Palearctic butterflies, a list
which is remarkably short in the western section of the Region.
Mr. Pocock kindly consented to test the hypothesis that J/elitcea
possesses the distasteful qualities of a model for mimicry, and
Commander J. J, Walker kindly helped to obtain material for

PALATABILITY OF SOME BRITISH INSECTS 867

the experiments which are recorded on pp. 825-827. The results
as a whole leave little doubt that Welitea is distasteful to many
birds, and that it does actually possess the qualities which would
render it an advantageous model for the Hesperiide.

Pages 827-830.

The experiments on Melanargia galathea are of peculiar interest.
The northern belt Satyrinz of this genus, with their white ground-
colour, stand out from the rest of their group. The under surfaces
are conspicuous, the species slow-flying and so abundant locally
that they may be called gregarious. The observations here
recorded show that they are also distinguished by their greater
distastefulness from other common northern Satyrines. The
appearance of the species of Melanargia, especially on the wing,
is markedly Pierine-like, and it is here also probable that a highly
distasteful genus has mimicked an assemblage of species which,
although generally less unpalatable, are excessively widespread and
abundant in individuals (see also p. 865).

Pages 830-831.

The experiments upon Lyczenide, Nemeobiine, and Hespertide
were not sufficiently numerous to form the foundation for safe
conclusions.

Pages 831-855.

The remaining experiments, for the most part, afford valuable
confirmation of previous work, but they also raise new questions
of great interest. Confirmation is afforded by the evident un-
palatability of Zygena, Hucheha, and Abrawas among the moths,
of the Saw-fly larve, of the Telephoride, Phytophaga, and Cocci-
nellidee among the beetles, and of the Hemiptera, as also by the
special and peculiar defensive secretions of the Carabide and, in
sharp contrast to all these observations, by the palatability of the
procryptically coloured moths and larvee.

Pages 847-848.

The apparent distastefulness of the humble-bee-like Volucella
bombylans suggests conclusions of so much importance and interest
that abundant confirmation is essential, and should be readily
available with so common a species.

Pages 848-852.

Experimental evidence that the Aculeate Hymenoptera possess
some special defence independently of the stings of the females is
now obtained for the first time. It was suggested as probable by
the present writer in 1904, as a result of the observation that the
males of the bee Sphecodes emerge in immense numbers and
form complex mimetic associations, before the appearance of the
females, as also from the consideration that the Braconide are
extensively mimicked *.

* Trans. Ent. Soc. Lond. 1904, pp. 645-6.

868 ON THE PALATABILITY OF SOME BRITISH INSECTS.

Page 849.

Although so many insectivorous animals in confinement dis-
regarded the special defence of Yormica rufa, there can be little
doubt that such defence is very effective in the wild state. It is
impossible on any other hypothesis to account for the conditions
under which the species exists—swarming in vast numbers in
restricted areas and an easy prey to any enemy that would dare
to attack.

A very important conclusion is suggested by several of the
experiments recorded in this memoir, namely, that the tastes of
mammals and birds are widely different. The author points out
that the defence of the ground-beetles appealed more strongly
to the mammals than to the birds, but it was also apparent in
many of the experiments that the unpalatability of conspicuous
Lepidoptera was, conversely, far more obvious to the birds than
to the mammals. Jn view of the part which birds are believed
to play in the production of mimetic resemblance, it is obvious
that this inference may be highly significant.

ADDENDUM.

Dr. P. Caatmers Mrrcneni’s Memoir ‘On Longevity and
Relative Viability in Mammals and Birds,” P. Z.8.
1911, p. 425.

[The Rev. F. C. R. Jourpatn has kindly called my attention to
the fact that I have overlooked a valuable paper “‘On the
Duration of Life of the Animals in the Zoological Garden at
Frankfort-on-the-Main,” by Director Dr. Max Scumunpt, P. Z.8.
1880, p. 299, and containing many valuable figures as to
longevity|.—P. C. M., Aug. 2, 1911.

No. 98.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.*
June 13th, 1911.

Epwin T. Newton, Esq., F.R.S., in the Chair.

The Minutes of the last Scientific Meeting were confirmed.

Mr. H. G. Purmmer, F.R.8., Pathologist to the Society, pre-
sented a Report on the Pathological Examination of Rats caught
in the Regent’s Park and in the Society’s Gardens. 500 rats
had been examined between the lst of January and the 17th of
May, 1911, all in a precisely similar manner. The spleen, lungs,
glands, and blood were examined microscopically; and from
any animal which looked in any way unhealthy cultures were
made.

The results were summarized as follows :—5 rats were caught
in the Park, and 495 in the Gardens: 283 of these were males
and 217 females.

3 rats had tubercle, 10 had tapeworm cysts in the liver, 49 had
Trypanosoma lewisi in their blood, 2 had empyema (not tubercular),
1 had a tumour of the lower jaw (the result of an old injury), and
1 had pleuritis and hydrothorax (not tubercular).

Bacteria were found in 71 rats: in 40 in the lungs, and in 31
in the spleen.

Saccharomycetes were found in the lungs of 16 rats.

Fleas were found on 4 rats, and lice on 8 rats.

The general condition of the rats was very good, and in none
was anything at all suspicious found.

* This Abstract is published by the Society at its offices, Zoological Gardens,
Regent's Park, N.W., on the Tuesday following the date of Meeting to which
it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge,
to all Fellows who subscribe to the Publications; but it may be obtained on the
day of publication at the price of Sixpence, or, if desired, sent post-free for
the sum of Six Shillings per annum, payable in advance.

B)
2)

Mr. R. I. Pocock, F.R.S., ¥F.Z.S., the Society’s Curator of
Mammals, exhibited the skin and skull of a Crested Rat
(Lophiomys ibeanus), sent from British Kast Africa by Mr. R. B.
Woosnam, and after drawing attention to the characters of the
skull in this genus, said that in his opinion the coloration was
for the purpose of self-advertisement and rendered the animal
conspicuous at night.

Dr. R. W. SHuFELDT, C.M.Z.S., sent for exhibition a photograph
he had taken of a living specimen of a male albino Woodchuck,
Arctomys monax, that had been sent to him from Virginia,

WESssA

Mr. R. E. Houpine exhibited and made remarks upon the
Horns of a Highland Ram, a Fallow Deer, and a Roebuck, which
were fused at the base, and also the skull of a coursing Greyhound
with abnormal dentition.

Dr. R. E. Draks-Brocxkmay, F.Z.8., read a paper on Antelopes
of the genera Madoqua and Rhynchotragus found in Somaliland,
specimens of most of which were exhibited at the Meeting. He
made general remarks on all the Dik-diks and gave a short
account of the species and subspecies, including the description
of a new form.

The Hon. Paut A. Meruuen, F.Z.8., communicated a paper
“On an Amphipod from the Transvaal,” in which he gave a
detailed description of a new freshwater Gammarid of the genus
Eucrangonyx, that had been found in caves in the Transvaal.

A paper was contributed by Mr. R. LypEKxKeEr on three African
animals. The first specimen was the skull of a Somali Rhinoceros,
a race for which the author adopted the name Rhinoceros bicornis
somalicus, Potocki. A Klipspringer skull from Northern Nigeria,
characterized by its great width and the peculiar form of the
lachrymal bone, was described as the type of a new race, Oreotragus
saltator porteust. Finally, a Gazelle from Algeria was referred
to a new species, Gazella hayi, agreeing approximately in size
with G. dorcas, but distinguished by the much straighter and
non-lyrate horns, each of which carried only about a dozen rings.
The face-markings were approximate to those of G. cuvierd.

A paper entitled ‘A Contribution to the Ornithology of Western
Colombia,” by Mr. C. E. HeLrtmayr, was communicated by Dr. P.
L. Scuarer, F.R.S. This memoir was based on a collection made
by Mr. M. G. Palmer in 1908 and 1909, which, though numbering
hardly 700 specimens, was of considerable interest and contained
many rare species, and also furnished information of importance
to students of zoogeographical problems.

©

No. 98.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.*
June 13th, 1911.

Epwin T. Newton, Esq., F.R.S., in the Chair.

The Minutes of the last Scientifie Meeting were confirmed.

Mr. H. G, Pumuer, F.R.S., Pathologist to the Society, pre-
sented a Report on the Pathological Examination of Rats caught
in the Regent’s Park and in the Society’s Gardens. 500 rats
had been examined between the Ist of January and the 17th of
May, 1911, all in a precisely similar manner. The spleen, lungs,
glands, and blood were examined microscopically; and from
any animal which looked in any way unhealthy cultures were
made.

The results were summarized as follows :—5 rats were caught
in the Park, and 495 in the Gardens: 283 of these were males
and 217 females.

3 rats had tubercle, 10 had tapeworm cysts in the liver, 49 had
Trypanosoma lewisi in their blood, 2 had empyema(not tubercular),
1 had a tumour of the lower jaw (the result of an old injury), and
1 had pleuritis and hydrothorax (not tubercular).

Bacteria were found in 71 rats: in 40 in the lungs, and in 31
in the spleen.

Saccharomycetes were found in the lungs of 16 rats.

Fleas were found on 4 rats, and lice on 3 rats.

The general condition of the rats was very good, and in none
was anything at all suspicious found.

* This Abstract is published by the Society at its offices, Zoological Gardens,
Regent's Park, N.W., on the Tuesday following the date of Meeting to which
it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge,
to all Fellows who subscribe to the Publications; but it may be obtained on the
day of publication at the price of Sixpence, or, if desired, sent post-free for
the sum of Six Shillings per annum, payable in advance,

38

Mr. R. I. Pococx, F.R.S., F.Z.8., the Society’s Curator of
Mammals, exhibited the skin and skull of a Crested Rat
(Lophiomys ibeanus), sent from British East Africa by Mr. R. B.
Woosnam, and after drawing attention to the characters of the
skull in this genus, said that in his opinion the coloration was
for the purpose of self-advertisement and rendered the animal
conspicuous at night.

Dr. R. W. Suure.pt, C.M.Z.8., sent for exhibition a photograph
he had taken of a living specimen of a male albino Woodchuck,
Arctomys monax, that had been sent to him from Virginia,

U.S.A.

Mr. R. H. Hoipine exhibited and made remarks upon the
Horns of a Highland Ram, a Fallow Deer, and a Roebuck, which
were fused at the base, and also the skull of a coursing Greyhound
with abnormal dentition.

Dr. R. EH. Draxs-Brocremay, F.Z.8., read a paper on Antelopes
of the genera Madoqua and Rhynchotragus found in Somaliland,
specimens of most of which were exhibited at the Meeting. He
made general remarks on all the Dik-diks and gave a short
account of the species and subspecies, including the description
of a new form.

The Hon. Paut A. Meruuen, F.Z.8., communicated a paper
“On an Amphipod from the Transvaal,” in which he gave a
detailed description of a new freshwater Gammarid of the genus
Hucrangonyx, that had been found in caves in the Transvaal.

A paper was contributed by Mr. R. LyDEKKER on three African
animals. The first specimen was the skull of a Somali Rhinoceros,
a race for which the author adopted the name Phinoceros bicornis
somalicus, Potocki. A Klipspringer skull from Northern Nigeria,
characterized by its great width and the peculiar form of the
lachrymal bone, was described as the type of a new race, Oreotragus
saltator porteusi. Finally, a Gazelle from Algeria was referred
to a new species, Gazella hayi, agreeing approximately in size
with G. dorcas, but distinguished by the much straighter and
non-lyrate horns, each of which carried only about a dozen rings.
The face-markings were approximate to those of G. cuvieri.

A paper entitled ‘‘ A Contribution to the Ornithology of Western
Colombia,” by Mr. C. EK. HELLMAYR, was communicated by Dr. P.
L. Scuater, F.R.S. This memoir was based on a collection made
by Mr. M. G. Palmer in 1908 and 1909, which, though numbering
hardly 700 specimens, was of considerable interest and contained
many rare species, and also furnished information of importance
to students of zoogeographical problems.

No. 99.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.*

June 27th, 1911.

FREDERIOK GiLLeTt, Esq., Vice-President,
in the Chair.

The Minutes of the last Scientific Meeting were confirmed.

The Secretary read a Report on the Additions that had been
made to the Society's Menagerie during the month of May
1911.

Dr. W. T. Caumay, F.Z.S., exhibited a number of living
specimens of the Brine Shrimp (Artemia salina) which had been
bred from Tidman’s Sea Salt. He remarked that this sea salt, as
sold in the shops, was found frequently to contain living eggs
of Artemia and that it was easy to obtain a supply of living
specimens. An 8 per cent. solution, allowed to stand for a few
days, produced a swarm of larve, and these could be fed on the
strained juice of green leaves and raised to maturity,

Mr. J. Lewis Bonunorz, M.A., F.Z.S., exhibited a pair of
Egyptian Desert-Mice (/eriones crassus) which showed a darker
and more rufous colour than normal examples. This coloration
had been artificially produced by keeping the animals in a moist
atmosphere at 80° Fahr. They were first exposed to these con-
ditions on the 7th of April and a month later were conspicuously
darker ; after that the darkening process still continued, but more
slowly, and they appeared for some time previous to being killed

* This Abstract is published by the Society at its offices, Zoological Gardens,
Regent’s Park, N.W., on the Tuesday following the date of Meeting to which
it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge,
to all Fellows who subscribe to the Publications ; but it may be obtained on the
day of publication at the price of Sixpence, or, if desired, sent post-free for
the sum of Six Shillings per annum, payable in advance,

A2

to have reached a limit to their darkening. During this same
period, and owing to the fine weather, other examples had been in
a temperature that rose during the day to 90°, falling at night to
60° or even lower. This had apparently produced no change in
their coloration. The change in the examples exhibited was
therefore due probably rather to the humidity than to the tempe-
rature. Mr. Bonhote was therefore inclined to think that the
pale colour of desert animals was due to the extreme dryness of
the atmosphere rather than to any special assimilation of their
colour to the surroundings.

Mr. R. I. Pocock, F.R.S., F.Z.8., Superintendent of the
Gardens, exhibited photographs of a hybrid between the Somali-
land Wild Donkey (Zquus asinus somaliensis) § and the Mountain
Zebra (Equus zebra) 2, and remarked that although Mountain
Zebras and domestic Asses had been previously crossed, no hybrid
had hitherto been produced between this Zebra and the Somaliland
race of Hgwus asinus. The period of gestation was twelve months
and three weeks exactly. The foal showed much greater resem-
blance to Hqwus asinus than to LH. zebra, the body being practically
unbanded, except for the spinal and shoulder stripes. The stripes
on the legs extended as high as the level of the belly and were
broader and more numerous than in the sire and narrower and
fewer than in the dam. In the presence of the shoulder stripe,
the spinal stripe, and the stripe on the base of the ear, the foal
resembled typical examples of Hquwus asiwus and differed from its
actual sire, which was without those marks.

Mr. D. Seru-Surru, F.Z.8., the Society’s Curator of Birds,
exhibited two immature Black-backed Porphyrios (Porphyrio
melanonotus) which had been bred in the Gardens, and remarked
upon their possession of a well-developed claw on the pollex.
Although these wing-claws were said to be functional only in the
Hoatzin amongst living birds, the exhibitor believed that they
were so also in the present species as also probably in the Common
Moorhen, these birds using these appendages to assist them in
climbing amongst reeds and other herbage.

The SECRETARY remarked that on a recent visit to the Ostrich
Farm of Mr. Carl Hagenbeck at Stellingen, near Hamburg, he
had seen in the incubator fertile eggs of Struthio massaicus from
German East Africa, S. australis from South Africa, and S. mo-
lybdophanes from Somaliland, the eggs all having been laid at
Stellingen. A. Reichenow (‘Die Vogel Africas,’ vol. i. p. 7) had
already described and figured certain specific differences in the
number and arrangement of the pits on the eggs of these species.
He himself had been interested to notice that the eggs of the
Masai Ostrich were much larger than those of the others, more
spherical in shape, and very smooth and porcelanous in texture.
Those of the Cape Ostrich were somewhat similar in shape and

43

texture, but were much smaller; Mr. Hagenbeck had informed
him that a pair of the Masai Ostrich bred by himself and sent out
to the Cape were regarded by expert ostrich farmers there as
unusually large birds. The eggs of the Somali Ostrich were
larger than those of the Cape Ostrich, but smaller than those of
the Masai species, and were markedly oval in shape with a rougher,
less polished surface.

The SzcrETARyY remarked that on his recent visit to Mr. Hagen-
beck’s Zoological Park at Stellingen, near Hamburg, he had the
pleasure of seeing a fine young pair of the common African
Rhinoceros, obtained from British East Africa, the exact locality
being unknown. The male closely resembled the ordinary figures
and mounted examples of the species, in that the skin appeared
to be smoothly stretched over the sides of the body, but the ears
were fringed with long tufts of hair. The female, on the other
hand, had ne hair on the margin of the ears, and the general
external appearance was very different. At first sight it seemed
as if it were in very poor condition, the ribs standing out through
the skin, but closer inspection showed that in reality the skin
of the flanks was disposed in thick, permanent folds, arranged
roughly like ribs. Thinking it possible that these differences
might indicate the existence of distinct races of the Rhinoceros,
on returning to London he had at once examined the Society’s
own pair of examples of this species, both of which had come from
British East Africa, probably somewhere near Nairobi. The
female, purchased in 1906, had the ears unfringed with hair, like
those of Mr. Hagenbeck’s female, but the rib-folds on the skin
were no more than indicated, although there were very heavy
permanent folds round the neck. In the male, obtained in the
current year from Nairobi as part of the King’s African Collec-
tion, the ears were fringed with hair as in Mr. Hagenbeck’s male,
whilst the rib-like folds on the skin were extremelystrongly marked,
as in the case of Mr. Hagenbeck’s female. The presence or
absence of the marginal fringe on the ears was therefore probably
either an individually variable or a sexual character. In the
absence of knowledge of the exact provenance of all the four
examples, nothing could be said as to whether or no the presence
of the rib-like permanent folds on the body were racial. Their
existence, however, as well as the presence of the heavy fold round
the neck, showed that it was not correct to distinguish the Asiatic
Rhinoceroses from those of Africa by the presence in the former
of permanent skin-folds. The neck-fold was almost identical in
both, whilst, although they were differently arranged, deep body-
folds occurred in both.

Dr. A. Suita Woopwarp, F.R.S., V.P.Z.S., communicated a
paper by Dr. R. Broom, C.M.Z.S., entitled ‘“‘On some new South
African Permian Reptiles.’

44

Mr. F. EK. Bepparp, M.A., F.R.S., F.Z.S., Prosector to the
Society, read a paper ‘“‘On two new Genera of Cestodes from
Mammals,” based on specimens collected from animals in the
Society’s Gardens.

A paper was received from Miss Ruta Harrison, entitled
“Some Madreporaria from the Persian Gulf; with a Note on the
Memoir and some Further Notes on Pyrophyllia inflata by
Sypney J. Hickson, M.A., D.Sc., F.R.S., F.Z.8.” This memoir
dealt with a collection of corals made by Mr. F. W. Townsend,
the most interesting species obtained being P. inflata and Tre-
matotrochus zelandice, the latter of which was identical with the
specimens from Cook’s Straits, New Zealand, described as Cono-
cyathus zelandic by Prof. Martin Duncan. A new species of
Heterocyathus was described, and Professor Hickson appended a
note on the affinities of Pyrophyllia.

Mr. C. L. Bounencer, M.A., F.Z.S., contributed a paper “ On
Variation in the Medusa of Merisia lyonsi,” based on an exami-
nation of 400 specimens. Nearly 14 per cent. of these were
found to be abnormal, and to fall naturally into two well-marked
groups containing completely distinct phenomena. The author
discussed these separately and in detail, and stated that he knew
of no-form in which such a variety of abnormalities occurred as
in Merisia.

Mr. Cyrit CrossLanD, F.Z.8., presented two papers, entitled
“The Marginal Processes of Lamellibranch Shells” and ‘“‘ Warn-
ing Coloration in a Nudibranch Molluse and in a Chameleon.”
Asa pendent to the second of these Sir CoartEs Exzot, K.C.M.G.,
C.B., F.Z.S., contributed a paper on ‘‘Chromodorids from the
Red Sea collected and figured by Mx. Crossland,” containing an
account of three species of Chromodoris, which were noteworthy
as being varieties of known species or forms hitherto imperfectly
described.

This Meeting closes the Session 1910-1911. The next Meeting
of the Society for Scientific Business will be held on Tuesday,
October 24th, 1911, at half-past Hight o'clock p.m.

45
The following Papers have been received :—

1. A. D. Imus, D.Sc., B.A.

On some Collembola from India, Burma, and Ceylon, with a
Catalogue of the Oriental Species of the Order.

2. Prof. P. P. Susuxin, C.M.ZS.

Ontogenetical Transformations of the Bill in Ardea cinerea.

3. EK. P. Stepsine, F.L.S., F.Z.8.

Game Sanctuaries and Game Protection in India.

Communications intended for the Scientific Meetings suould
be addressed to

P. CHALMERS MITCHELL,

Secretary.

ZOOLOGICAL SoctEty OF LONDON,
Recent’s Park, Lonpon, N.W.
July 4th, 1911.

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Exhibitions and Notices (continued).

- Page
Mr. R. I. Pocock, F.B.S., F.L.S., F.Z.8. Exhibition of some hair from the “ puppy coat”

of a Grey Seal (Halicherus grypus) ......++++0s Rertiblonenan do ac Ghd 6G cmH OUIDO He 696

Mr. EB. G. Bounencer. Exhibition of living male specimens of the Midwife Toad (Alytes
obstetricams) carrying the eggs... 0. os ee nsw e se nsec ne wee cee nee e en aes 696

Mr. A. HE. Anpersoy. Exhibition of photographs of fossil mammals in the American
Museum of Natural History..............- Peongour Bhs taeee ez adaicfaievabareas b netieborage. eyerels 696

PAPERS.
26. On the Amphipod Genus Leptocheirus. By E. W. Sexton, Marine Biological Labora-

tory, Plymouth. (Plates XVII.XIX. and Text-fig. 146.) ....-.....-.-2..20--- «. 561
27. Notes on Marine Ostracoda from Madeira. By G. Srzwarpson Brapy, M.D., LL.D.,

D.Se., F.R.S., C.M.Z.8. (Plates XX.—XXIT.) 1.0... cw eee eee eee eee eee eee Toy 8B)
28. On Colour and Colour-pattern Inheritance in Pigeons. By J. Lewis Bonuors, M.A.,

F.LS., F.Z.8., and F. W. Swanzey, ¥.Z.8. (Plates XXIII.—-XXVI.) .............- 601

29. Contributions to the Anatomy and Systematic Arrangement of the Cestoidea. By
Frank E. Bepparp, M.A., F.R.S., F.Z.S., Prosector to the Society. (Text-figs. 148-159.) 626

30. On the Natural History of Whalebone Whales. By J. A. Morcu, Christiania.

(Bex totes NGO UG ao) tare yeteneteiavel afegovs) ia" 20 -ths (olal-e’e “a. ale eleveies @tietutey nieleiieieme/ ae fey tela eats 661
31. On Three New Trematodes from Reptiles. By Witttam Nicoun, M.A., D.Sc., M.B.,
TRASH (GPileiais O10 ey DO. \ AEE) eee Sion dis Gincin bn OS bdbboUona eb edbomooo ne uDeo eS 677

32. The Duke of Bedford’s Zoological Exploration of Eastern Asia —XIV. On Mammals
from Southern Shen-si, Central China. By Oxprietp Tuomas, F.R.S., F.Z.S.
(CRIB IG) 2 O81 DO) nin a Beer be Rae Eo odie goon ace NEUF aha tenets ofchiay ten tat aerate eee oka 687

33. An Investigation into the Validity of Miillerian and other forms of Mimicry, with
Special Reference tothe Islands of Bourbon, Mauritius, and Ceylon. By NrviniE

Manpurs, Lieut.-Colonel, R.A.M.C., B.Z.8., FLHLS...--. 000.0 cc cece ce cece eee cane 696
34. The Distribution of the Avian Genus Megapodius in the Pacific Islands. By J.J. Listur,
MinAcuRb Ns Huy Se ebeZ omen (BexhticUOOs)ie watelsttnts +) se inlelser state exe e/eraistol ain els, oraieiaiels 749

35. Contributions to the Morphology of the Group Neritacea of the Aspidobranch
Gastropods.—Part Ii. The Hznicinips. By Gitperr C. Bourng, M.A., D.Sc., E.RBS.,
IE Za e (a ates NCNONG == hc DB ete ter cinta’ st ney eaves feet cheval eiataial suelo natavatg eters (ch alens “harbors 759

36. On the Palatability of some British Insects, with Notes on the Significance of Mimetic
Resemblances. By R. I. Pocock, F.R.S., F.L.S., F.Z.S., Superintendent of the Society’s
Gardens and Curator of Mammals. With Notes upon the Experiments by Prof. EH. B.
POUTINONS HERES. HLA Sy tevepcrter a aicrase te srcral clei nist stsnclov ale eats’ eVatatstevelovarct Sercvere(el ccc aletel ateleine 809

ADDENDUM.

Additional reference to Dr. P. Cuatuers Mircnent’s memoir, “On Longevity and
Relative Viability in Mammals and Birds” ........-.....++.... SOM LEpeU aI erKCr 868

DESEO PLATES:
1911, Parr III. (pp. 557-868).

Plate Page

XVII. Leptocheirus pilosus Zaddach ......-.2+.....0----.-00-> )

XVIII. i-9. Leptocheirus guttatus Grube. 10-12. L. pinguis |
. Stimpson. 13-16. LZ. hirsutimanus Bate. 17-20. ae

EE bispinosuswNOnManeyas eet siete aera
XIX. Leptocheirus pectinatus Norman .......--+..++ee-eeeee }
XX.
XXI. } Ostracoda from Madeira ..........0e.ceee Obie a HeNAr 595
XXII. |
XXIIL. 1. Blue, 2. Silver, 3. Chequer Pigeons ................. -)
XXIV. 1. Dark Grizzle, 2 - White (crue 3. Grizzled Chequer | |
TEGO) | Jobadodssabpesccsos Wan soooscsducdgg ss G0 L601
XXV. 1. Dark Mealy, 2. Light Mealy, 3. White Mealy Pigeons “+i
XXVI. Feathers showing pattern-markings in Pigeons .........-
XXVII. Lechriorchis CHGDIT AN ING HA A AEO OG o UO aa Oaoe Oo teens \ 677
XXVIII. 6,7. Ochetosoma formosum. 8-10. Dasymetra conferta
XXIX. The Chinese Takin, Budorcas bedfordi ...- 1.0. ..00+ee0es 687
XXX. \
KAKI
XXXII. |
XXXIII. |
XXXIV. |
XXXYV. |
XXXVI. } Morphology of Ge WERPI CITES) sWo seit b taghte im date hts. iets 759
XXXVII. Fe
XXXVITII.
XXXIX. |
XL: '|
XI: |
XLII. )
NOTICE.

The ‘ Proceedings’ for the year are issued in fowr parts, paged consecutively,
so that the complete reference is now P. Z. 8. 1911, p.... The Distribution

is as follows :—
Part I. issued in March

PEA te East June.
srg ep eche E BlD a September.
EESRTUS EAS NO EE ak December.

‘ Proceedings,’ 1911, Part II. (pp. 181-555), were published on
July 6th, 1911.

‘The Abstracts of the Proceedings,’ Nos. 98 and 99,
are contained in this Part.

5
PROCEEDINGS

OF THE

GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

OF THE

ZOOLOGICAL SOCIETY
OF LONDON.

1911.

PART TV.

CONTAINING Paces 869 to 1213, witH 22 Prates
AND 64 TEXT-FIGUKES.

ae

| FEB 96 1919

DECEMBER 1911.

PRINTED FOR THE SOCIETY,
SOLD AT THEIR HOUSE IN REGENT’S PARK.
LONDON :

| MESSRS. LONGMANS, GREEN, AND CoO., ‘
| PATERNOSTER ROW. |

pT Ki sala alt
s Teri } Sots *

4
}
sy

Orn [Price Twelve Shillings. ] | ac

LSet Ol 0 IN: IN ans:

1911, Part IV. (pp. 869-1213).

EXHIBITIONS AND NOTICES.

{Pn Secretary. Report on the Additions to the Society’s Menagerie during the month
of April 1911

or gt PO ee OH OO PEE Eo ee ee EE ee Sr. sce ee er eens mrs OF eersteosce ee Our eeosae

The Rey. A. Mites Moss, M.A., F.Z.8S., F.E.S. Notice of a Memoir on the “‘ Sphingidze
of Peru ”

| os %) ex0) c|'e| e/'e\'8)@)/e a) 8) 8. 02 [sss 0) (sie ©) 6)'s, > s\» © e\ia: elie c/s le) e \eif/ip 0 ajc elle ee isvelie!a) jis) © mi elejieleialsl

Mr. H. G. Primer, F.R.S. Report on the Pathological Examination of Rats (Mus

decumanus) caught in the Regent's Park and in the Society's Gardens

Dr. R. W. Saurecpt, C.M.Z.S. Exhibition of a photograph of amale albino Woodchuck
(Arctomys moni)

Mr. R. E. Houpixe. Exhibition of horns of Deer and the skull of a Greyhound

ee eere

Mr. R. I. Pococn, F.R.S., 7.2.8. Exhibition of the skin and skull of the Crested Rat
(Lophiomys ibeanus Thos.), (Lext-fig. 190.)

i ee ee rc ee err rn

Tne Secretary. Report on the Additions to the Society’s Menagerie durine the month -
P y 8 g

of May 1911

CC a ie ee ek a i ir

Mr. D. Sern-Ssirno, F.Z.8. Exhibition of two immature Black-backed Porphyrios (Por-
phyrio melanonotus) with wing-claws. (Text-fig. 200.) .....-.......--

Mr. J. Lewis’ Bonnorts, M.A., F.L.S., ¥.Z.8. Exhibition ofa pair of abnormally coloured
Kgyptian Desert-Mice (Merzones crassus)

Ce i Ce Ci Ce rr ry

Dr. W. T. Catan, F.Z.S. Exbibition of living specimens of the Brine Shrimp (Artemia
salina)

SCC ee ee ee ee i a, i arid

Tne Secrerary. Remarks upon Ostrich eggs seen at Mr. Carl Hagenbeck’s Ostrich
Farm at Stellingen ......... Bass qososusc S66s0 06x SEA US SS suiao doo webs bo de dc

Tis Secretary. Remarks upona pair of young African Rhinoceros seen at Stellingen ..
g §

Mr. R. I. Pococs, F.R.S., F.Z.8. Exhibition of photographs of hybrid Zebra Feals.
(Dext-figs. 201=203))\ oe .e. oe 3 ee Bivens eee ndeleneasbatae Sinton choise Se eS eee ote

PAPERS.

37. The Alcyonaria of the Cape of Good Hope and Natal.—Goreonacna. By J. Srvant
Tuomson, Ph.D., F.R.S.E., F.L.8., Lecturer and Senior Demonstrator in Zoology,
University of Manchester. (Pls. XLIII.—XLV. and Text-fig. 167.) ...........

se

38. On the Structure of the Skull in Cynodont Reptiles. By R. Broom, M.D., D.Sc.,
OLIWIYAIS, (GHG IONE eral akesacltech IGS ASO)) Sodoscbdéodsaccos coodotgubbadoons

39. Tooth-germs in the Wallaby (Macrapus billardieri). By A. Hornweut-Ssitu,
L.R.C.P., M.R.C.S., and H. W. Marerr Tims, M.A.. M.D., F.LS., F.ZS.
(QE DOU \MULG eval Nexiairash NSW) Ak Gros sondodd0odooddnooosdno cous stosoiercnere

986

987
987

988

926

Contents continued on page 3 of Wrapper

THE ZOOLOGICAL SOCIETY OF LONDON,

eee

Tus Society was founded in 1826 by Sir Sramrorp RaFrtes,
Mr. J. Sasrnz, Mr. N. A. Vigors, and other eminent Naturalists,
for the advancement of Zoology and Animal Physiology, and for the
introduction of new and curious subjects of the Animal Kingdom,

and was incorporated by Royal Charter in 1829.

Patron.

HIS MAJESTY THE KING.

COUNCIL.
HIS GRACE THE DUKE OF BEDFORD, K.G., President.

THe Eart or ALTAMonrt, F.S.A.
Sir J. Rosp Braprorp, K.C.M.G.,

M.D:, DSe., F.R.S., Vice-
President.
Lr.-Cot. Srr R. Havetocx

Cuarues, M.D., K.C.V.O.
Atrrep H. Cocks, Esa., M.A.
Tue Rr. Hon. tar Earn or

Cromer, P.C.,G.C.B.,G.C.M.G.
F. G. Dawrrey Drewirrt, Esa.,

M.A., M.D.
Caartes DrumMonp,

Treasurer.

Sir Epwarp Douranp, Br., C.B.
Freperick Giiterr, Ese., Vice-
_ President.

Sipney F, Harmer, Ese., M.A.,

Sc.D., F.R.S., Vice-President.

Ksa.,

Srr Warter’ Roeper Lawrence,
Br., G.C.1.E.

Str Epmunp G. Loner, Br.

_E. G. B. Meave-Watpo, Esa.,
Vice-President.

P. Caatmers Mircuent, Ese.,
Mev Discs Ju Dever Rist.
Secretary.

| W. R. Ocitvie-Grant, Ese.
| Aprran D. W. Pottocg, Ese.

AvuBryn Trevor-Batrysg,
M.A.

AntHony H. WrneFrtetp, Esa.

A.Smira Woopwarp, Hse.,LL.D,
F.R.S., Vice-President.

Henry Woopwarp, Esea., LL.D.,
F.RS., Vice-President.

Ksa.,

9

vay

The Society consists of Fellows, and Honorary, Foreign, and
Corresponding Members, elected according to the By-Laws. It
carries out the objects of its foundation by means of the collection
of living animals, by its Library, and by its Scientific Publications.

The Office of the Society, Regent’s Park, N.W., where all com-
munications should be sent, addressed to “The Secretary,” is open
from Ten till Five, except on Saturdays, when it closes at Two p.m.

The Library, under the superintendence of Mr. F, H. Waterhouse,
is open daily at the above hours, except in September.

The Meetings of the Society for General Business are held in the
Meeting Room at the Society’s Office on the third Wednesday in
every month of the year, except in September and October, at half-
past Four o’clock p.m.

The Meetings for Scientific Business are held in the Meeting
Room at the Society’s Office fortnightly on Tuesdays, except in
July, August, September, and December and January, at half-past
Hight o’clock p.m.

The Anniversary Meeting is held on the 29th. of April, or the
nearest convenient day, at Four p.m.

The Society’s Gardens are open daily from Nine o’clock until
Sunset. Mr. R. I. Pocock, F.R.S., F.LS., is the resident Super-
intendent and Curator of Mammals, Mr. D. Seth-Smith is Curator
of Birds and Inspector of Works, and Mr. E. G Boulenger is:
Curator of Reptiles. The Prosectorium for Anatomical and Patho-
logical work is under the charge of Mr. Frank E. Beddard, M.A.,
F.R.S., Prosector, assisted by Mr. H. G. Plimmer, I.R.S., M.R.C.S.,
Pathologist to the Society.

TERMS FOR THE ADMISSION OF FELLOWS.

Frriows pay an Admission Fee of £5, and an Annual Contri-
bution of £38, due on the Ist. of January, and payable in advance,
or a Composition of £45 in lieu thereof; the whole payment,
including the Admission Ice, being £50.

No person can become a Friiow until the Admission Fee and
first Annual Subscription have been paid, or the annual payments
have been compounded for.

Fettows elected in November and December are not liable for
the Subscription for the current vear.

3
PRIVILEGES OF FELLOWS.

Ferttows have Personal Admission to the Gardens upon signing
their names in the book at the entrance gate, and may introduce
Two Companions daily.

The Wire or Hussanp of a Fattow can exercise these privileges
in the absence of the Fellow.

livery Frtiow is entitled to receive annually 60 undated Green
Cards, and, when no specific instructions are received, the supply
will be sent in this form. If preferred, however, 20 Green Cards
may be exchanged for a book containing 2 Orders for each
Saturday * throughout the year. A similar book of Sunday Orders
may also be obtained in lieu of 20 Green Cards. A Green Card
may also be exchanged for 2 Butf Cards for the use of Children
under 12 years of age.

It is particularly requested that Fellows will sign every Ticket
before it goes out of their possession, Unsigned ‘Tickets are not
available.

Green and Buff Tickets may be used on any day and in any year,
but in no case can two Children be admitted with one Adult's
Ticket, or an Adult be admitted with two Children’s Tickets.

The annual supply of Tickets will be sent to each Frtrow on the
Ist. of January in every year, upon filling up and returning the form
of Standing Order supplied to Fellows.

Frttows are not allowed to pass in friends on their written
order or on presentation of their visiting cards.

Futtows are exempt from payment of the fee for Painting,
Sketching, and Photographing in the Society’s Gardens.

Frttows have the privilege of receiving the Society’s ordinary
Publications issued during the year upon payment of the additional
Subscription of One Guinea. This Subscription is due upon the
Ist. of January, and must be paid before the day of the Anniversary
Meeting, after which the privilege lapses. Frtiows are likewise
entitled to purchase these Publications at 25 per ceut. less than
the price charged to the public. A further reduction of 25 per
cent. is also made upon all purchases of Publications issued prior
to 1881, if above the value of Five Pounds.

Frrttows also have the privilege of subscribing to the Annual
Volume of ‘The Zoological Record,’ which gives a list of the Works
and Publications relating to Zoology in each year, for the sum of

* The Saturday Orders are not available if the Fellow introduces friends
personally on that day.

4

One Pound Ten Shillings. Separate divisions of volumes 39 to
42 can also be supplied. Full particulars of these publications can
be had on application to the Secretary.

Frntows may obtain a TransrerasLe Ivory Ticker admitting
two persons, available throughout the whole period of Fellowship,
on payment of Ten Pounds in one sum, A second similar ticket
may be obtained on payment of a further sum of Twenty Pounds.

Any Frttow who intends to be absent from the United Kingdom
Curing the space of at least one year, may, upon giving to the
Secretary notice in writing, have his or her name placed upon the
“dormant list,” and will then be called upon to pay an annual
subscription of £1 only during such absence, but after three years
must make a further application to be retained on that list.

Any Frtiow, having paid all fees due to the Society, is at liberty
to withdraw his or her name upon giving notice in writing to the
Secretary.

Ladies or Gentlemen wishing to become Fellows of the Society

are requested to communicate with the undersigned.

P. CHALMERS MITCHELL,

Secretary.
Regent’s Park, London, N.W.,
December, 1911.
MEETINGS
OF THE
ZOOLOGICAL SOCIETY OF LONDON
FOR
SGILIB IN POI 1G) 1s) WES ICIS! IBS) Si
1912.
Truspay, Fepruary.. 6 & 20 | Tunspay, JuNF...... 4
55 INRCEH 2a) pOROg ee 3 OcroBER .. 29
“4 NPR See yOu Ze Ee November... 12 & 26
IMA ee 2k 7 & 21

The Chair will be taken at half-past [ight o'clock in the Evening
precisely.

ZOOLOGICAL SOCIETY OF LONDON.

LIST OF PUBLICATIONS.
Tue scientific publications of the Zoological Society of London
are of two kinds—“ Proceedings,” published in an octavo
form, and ‘‘ Transactions,” in quarto.

According to the present arrangements, the “ Proceedings”’
contain not only notices of all business transacted at the scien-
tific meetings, but also all the papers read at such meetings
and recommended to be published in the ‘‘ Proceedings ”’ by
the Committee of Publication. A large number of coloured
plates and engravings are issued in the volumes of the
“ Proceedings,” to illustrate the new or otherwise remark-
able species of animals described therein. Amongst such
illustrations, figures of the new or rare species acquired in a
living state for the Society’s Gardens are often given.

The “Proceedings” for each year are issued in four
parts, paged consecutively, in the months of March, June,
September, and December. From January 1901 they have
been issued as two half-yearly volumes, indexed separately.

An “ Abstract of the Proceedings” is published by the
Society on the Tuesday following the date of Meeting to
which it refers. It is issued along with the “ Proceedings,”
free of extra charge, to all Fellows who subscribe to the
Publications, but it may be obtained on the day of pubii-
cation at the price of Sixpence, or, if desired, sent post free
for the sum of Six Shillings per annum, payable in advance.

The “Transactions ”’ contain such of the communications
made to the scientific meetings of the Society as, on account of
the nature of the plates required to illustrate them, are better
adapted for publication in the quarto form. They are issued
at irregular intervals.

Fellows and Corresponding Members, upon payment of
a Subscription of One Guinea before the day of the Anni-
versary Meeting in each year, are entitled to receive the
Society’s Publications for the year. They are likewise
entitled to purchase the Publications of the Society at 26 per
cent. less than the price charged for them to the Public. A
further reduction of 23 per cent. is made upon purchases of
Publications issued prior to 1881, if they exceed the value of
five pounds.

Fellows also have the privilege of subscribing to the
Annual Volume of the Zoological Record for a sum of 30s.
(which includes cost of delivery), payable on the Ist. of July
in each year; but this privilege is forfeited unless the
subscription be paid before the 1st. of December following.

The following is a complete list of the publications of the
Society already issued.

TRANSACTIONS* OF THE ZOOLOGICAL SOCIETY OF LONDON.

4to. 19 vols. and Index. Eicetto Price tote
ellows. ublie.

Vol. I., containing 59 Plates.... (1833-35) .... £3 138 6 .... £418. OF
A - a Wt pease CISSO=4N) ea. Wale sO pmOne @ (oi
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Eh HNGLENG, ty DAEs (COS SNSTO) ROMA TsO ep reeloielonene

PROCEEDINGS OF THE COMMITTEE OF SCIENCE AND
CORRESPONDENCE OF THE ZOOLOGICAL SOCIETY OF

LONDON. 8vo. 2 vols. (Letterpress only). 22 ' Pace
Pave Me ISOS ILO G@s “Gesaooccagocas AS Gs) ee BOS
4 Ue TEED. Ean ene Ms Gi os

PROCEEDINGS OF THE ZOOLOGICAL SOCIETY OF LONDON.
8vo. 15 vols. (Letterpress only) and Index. (First Series.)

: Price to Price to the Price to Price to the

Fellows. Public. Fellows. Public.

Part J. 1833. 1 vol. 8vo. 4s. 6d. .. Gs.f | Part IX. 1841.1 vol. 8vo. 4s. 6d. .. 68.F
Pe Wieeade = ABS ag (Gp be X. 1842. ke ASG. meen Ose

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ViIL. 1840. . ° 4s.6d... 6st | Index 1880-1847. , 4s.6d. .. 6s.

aN oe
8vo. 18 vols. and Index. (Second Series.)
Letterpress only. With Plates coloured.
Price to Price to the Price to Price to the
Fellows. Public, Fellows. Public.
Part SWAT, WSS. Ul swwoll, Bras 4's, Ge 45 CS. ooscaocen £10058) 6 SIG
25 XVI. 1849. 5 AS ROG ancy OSI ee ciate tence 1 (0. 68 ooo) DGG
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Index 1848-1860. . ds. 6d. 6s,

t+ Out of print.
* In consequence of a re-arrangement of the stock of the ‘Transactions,’ the Society is
now able to offer for sale, at the reduced price of £30, sets of Vols. v.-xvi. inclusive, and
separate papers, of which a list can be supplied, at about one-fourth their published price.

PROCEEDINGS OF THE SCIENTIFIC MEETINGS OF THE
ZOOLOGICAL SOCIETY OF LONDON. 8vo. 40 vols. and 4 Indices.

Letterpress only. With Plates uncoloured. With Plates coloured.

Price to Price to the Price to Price to the Price to Price to the

Fellows. Public. Fellows. Public. Fellows. Public.
ISIE: 5g 2b Othivco5.6 OSs oomebc 9s, DEE LOS erate’ BRe, Oh sooo GaRaT
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* No perfect copies in stock. ft Out of print.

PROCEEDINGS or roe GENERAL MEETINGS ror SCIENTIFIC
BUSINESS or tar ZOOLOGICAL SOCIETY OF LONDON.

8vo. 22 vols.

Price to Price to the
Fellows. Public.
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LISTS OF THE ANIMALS IN THE SOCIETY’S GARDENS.

List of the Vertebrated Animals now or lately Living in the Gardens
of the Zoological Society of London. (Mighth Edition.) 8vo.
1883. Cloth, 4s. 6c.

List of the Vertebrated Animals now or lately Living in the Gardens
of the Zoological Society of London. (Ninth Kdition.) 8vo.
1896. Cloth, 6s.

Catalogue of the Library of the Zoological Society of London
(Fifth Edition.) Svo. 1902. Cloth, 6s.

THE OFFICIAL ILLUSTRATED GARDEN GUIDE— 9th Edition
—with (1) a Railway and ‘Street Map, showing a direct
route to the “* Zoo” from all parts of London and the Suburbs ;
(2) a Plan of the Grounds, showing at a glance the location of
the animals; (3) a short description of some of the principal
animals in the Collection (now containing over 3000 spe-
cimens), together with 50 Photographic Illustrations and
Index. Price 6d. in Stiff Paper Cover, postage 13d., or in
Green Cloth Cover price 1s. 2d. post free.

P. CHALMERS. MITCHELL,
Secretary.
Regent's Park, London, N.W.,
December, 1911.

These publications may be obtained at the Socrrry’s Orrice,
at Messrs. Lonemans (Paternoster How, £.C.), or through any
bookseller.

AVOLOGICAL SOCIETY OF LONDON.

THE ZOOLOGICAL RECORD.

HE object of the Zoonoeican Kecorp is to give, by means of an
annual Volume, complete lists of the Works and Publications
relating to Zoology in all its branches that have appeared during
the year preceding the issue of the Volume, together with full
information as to the points they deal with, arranged in such a
manner as to serve as an Index to the literature of Zoology in all
parts of the globe, and thus to form a repertory that will retain its
value for the Student in future years.

The ‘ Zoological Record’ having been amalgamated with the
International Catalogue of Scientific Literature, Zoology, Volumes
from 43 onwards cam now be obtained only from Messrs. Harrison
& Sons, except when purchasing complete sets from the Zoological
Society.

Under the scheme of amalgamation, Fellows of the Society, and
Institutions already on the subscription-list, have the privilege of
subscribing at the old rate of 30s. per annum, which covers the
cost of carriage of the volume. The subscription becomes due on
July Ist. in each year, and lapses if not paid by the 1st. of December
following.

The Society is able to supply complete sets of the Record on the
following terms :—

Vols. 1 to 37, price £14 10s. net.
Vols. 38 to 42 at 10s. each net.
Vol. 43 and onwards at 40s. each.
The prices for separate volumes ere as follows :-—
Vols. 1 to 42 (except Vols. 4 and 6) 10s. each net.
The price of the ‘Zoological Record,’ Vol. 43 and subsequent volumes,
published now by Messrs. Harrison and Co., is 40s. each.

Inpex Zootocicus. An alphabetical list of names of genera
and subgenera proposed for use in Zoology, as recorded in the
‘Zoological Record,’ 1880-1900; together with other names not
included in the ‘ Nomenclator Zoologicus’ of S. H. Scudder. Com-
piled (for the Zoological Society of London) by CHarrus Ownn
Wartrrnouse and edited by Davin Sarr, Editor of the ‘ Zoological
Record.’ London, 1902. Price to Fellows, 18s.; price to the
public, 20s., or if sold with a set, 10s.

Divisions of the ‘ Zoological Record’ of Vols. 39 to 42 ean be
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y Messrs.

P. CHALMERS MITCHELL,
Secretary.

Recents Park, Lonpon, N.W.
December, \911.

ON ADDITIONS TO THE MENAGERIE. 869

EXHIBITIONS AND NOTICES.
May 23, 1911.

Dr. A. Smrra Woopwarp, F.R.S., Vice-President,
in the Chair.

THE Secrerary read the following report on the additions made
to the Society’s Menagerie during the month of April, 1911 :—

The number of registered additions to the Society’s Menagerie
during the month of April last was 274. Of these 75 were
acquired by presentation, 147 by purchase, 34 were received on
deposit, 5 in exchange, and 13 were born in the Gardens.

The number of departures during the same period, by deaths
and removals, was 169.

Amongst the additions special attention may be directed to :—

1 Black Serval (Felis serval), from Nairobi, presented by the
Marchioness of Donegal to H.M. Tue Krne’s African Collection,
on April 25th.

2 Bactrian Camels (Camelus bactrianus), from Central Asia, and
1 Yak (Poéphagus grunniens), from Tibet, presented on April 4th
and 29th respectively, by H.G. The Duke of Bedford, K.G.,
President of the Society.

1 Klipspringer ( Orectragus saltator), from Somaliland, presented
by R. E. Drake-Brockman, Esq., M.R.C.S., F.Z.S., to H.M. Tax
Kine’s African Collection, on April 28th.

1 Savannah Deer (Dorcelaphus savannarum), from Iquique,
Chile, presented on April 24th by Miss Peggy Lomax,

1 Black-tailed Oribi (Ourebia nigricaudata), from the Ivory
Coast, presented by F. W. F. Jackson, Esq., to H.M. Tue Kina’s
African Collection, on April 4th.

2 Australian Bee-eaters (Jerops ornatus), from New South
Wales, new to the Collection, purchased on April 30th.

2 Woodhouse’s Jays (Aphelocoma woodhousei), from North
America, and 3 Pigmy Ground-Doves (Chamepelia minuta), from
South America, new to the Collection, received in exchange on
April Ist.

2 Sharpe’s Wood-Owls (Syrnium nuchale), 1 Fraser’s Eagle-Owl
(Bubo poénsis), | Beautiful Wood-Hawk (Dryotriorchis spectabilis),
and 1 One-streaked Hawk (Melierax monogrammicus), from
Dunkwa, Gold Coast, presented by Dr. H.G. F. Spurrell to
H.M. Tae Kine’s African Collection, on April 2nd.

4 Gigantic Salamanders (Megalobatrachus maximus), from Japan,
purchased on April 28th.

The Rev. A. Mines Moss, M.A., F.Z.S., F.E.S., gave a short
account of his memoir on the Sphingide of Peru, based on studies
of Lepidoptera, with special reference to the larvee, which he had
made during a three years’ residence at Lima.

This paper will be published in the ‘Transactions’ in due
course.

Proc. Zoou. Soc.—1911, No. LX. 60

870 DR. J. STUART THOMSON ON

PAPERS.

37. The Aleyonaria of the Cape of Good Hope and Natal.—
GorconaceaA. By J. Stuart THomson, Ph.D., F.R.S.E.,
F.L.S., Lecturer and Senior Demonstrator in Zoology,
University of Manchester *.

[Received May 4, 1911: Read May 23, 1911.]

(Plates XLIII.—XLV.? and Text-figure 167.)

The Aleyonaria here reported on were collected during the years
1898 to 1907, off the coasts of Cape Colony and Natal. In
a previous paper I dealt with the Alcyonacea, and in the
present report I confine myself to the Gorgonacea. This con-
tribution is not, however, to be regarded as an exhaustive one, as,
unfortunately, the specimens have come into my hands at different
times. Professor Hickson has contributed two papers on South
African Aleyonaria to the publication, ‘ Marine Investigations in
South Africa,’ and I have to thank him for allowing me to
examine his type-specimens and microscopic slides of the spicules
of various forms.

I have also to thank Professor J. D. F. Gilchrist, formerly
Government Biologist at the Cape of Good Hope, who kindly
entrusted these Aleyonarians to me for description.

Tn this paper, the following 19 species are described or recorded,
of which 6 are new :—

Family BRIAREIDA.

Suberia capensis, sp. n., p. 871.
Spongioderma verrucosum Mobius, p. 874.

Family MEurropip#.

Melitodes esperti W. & S., p. 874.
Melitodes nodosa W. & S., p. 876.
Melitodes dichotoma Pall., p. 877.

Family Istpm.
Ceratoisis ramosa Hickson, p. 877,

Family Muriceis.

Muriceides fusca, sp. n., p. 878.
Acanthogorgia armata Verrill, p. 880.

* Communicated by Prof. ARTHUR DENpDy, D.Sc., F.R.S., F.Z.S.

+ For explanation of the Plates see pp. 892-893.

+ “The Alcyonaria of the Cape of Good Hope and Natal.—Alcyonacea” (with
our plates). ‘Trans. Roy. Society Edinburgh, vol. xlvii. part ii. (No, 19), 1910,

P.Z.S. 1911. Pl. XLII.

S
.%
= VS
fi “ * e
ad
x
ae
4 ,
Ree
I ¢ 3
Soy! :
id
Sy &
+o
~ s .
e 3% vip ey

ALCYONARIA OF

THE CAPE OF GOOD HOPE.

P.Z.S8.1911.P1. XLIV.

ALCYONARIA OF THE CAPE OF GOOD HOPE.

P.Z.8.1911.P1. XLV.

CAPE OF GOOD HOPE.

ALCYONARIA OF THE

CORALS FROM SOUTH AFRICA. 871

Family PLEXAURIDS.

EKunicella papillosa Esper, p. 880.
Psammogorgia pulchra, sp. n., p. 881.
Huplexaura media, sp. n., p. 883.

Family MALAcocoreiiD&.
Malacogorgia capensis Hickson, p. 884.

Family Primnoms.

Stachyodes gilchristi, sp. n., p. 885.
Thouarella hicksoni, sp. n., p. 886.

Family GorGoNntD&.

Gorgonia capensis Hickson, p. 887,
Gorgonia flammea HK. & S., p. 888.

Family GoRGONELLID#.

Scirpearia furcata emend. Simpson, p. 888.

Scirpearia flagellum emend. Simpson, p. 888.

Hicksonella spiralis Simpson = Juncella spiralis
Hickson, p. 889.

Family BRIAREIDS&.

SUBERIA CAPENSIS, sp. n, (Pl. XIII. fig, 4; Pl. XLV.
figs. 4a—c.)

The colony is of a dark red colour, resembling that of Gorgonia
flammea K. & 8.,and consists of (1) an encrusting base, (2) a more
or less cylindrical trunk, and (3) an expanded apical part. The
basal part is 10 mm. long, 13 mm. broad, and 6 mm. high,
The trunk is about 27 mm. in height, 6 mm. in diameter basally
and medially, and about 7 mm. apically. The lowest polyp is
situated on the trunk about 13 mm. from the base. About
fourteen polyps are situated near the apex of the trunk, and these
ave almost entirely confined to three sides. The largest polyps of
this part are 2 mm. in height and 3 mm. in diameter. The
distance of the polyps from one another varies considerably,
nainely from about 1 to 8 mm. The apical part of the colony is
about 14 mm, in height and 15-17 mm. in diameter (including
the polyps). The polyps are in this part more prominent than
in the trunk region. They vary from 2-3 mm. in height and
breadth. The polyps are irregularly scattered over the surface of
the apical part, and the interval between them varies from about
1to5mm. There are thirty to forty polyps in the apical part.

The entire surface of the colony is rough to the touch and has
a tough consistency. In the centre of the colony there is a horny
and limy axis; in other words, the axis consists of horny tissue
and in association with this a large number of spicules. The
spicules of the axis usually differ in form from those of the cortex.

60*

872 DR. J, SEUART THOMSON ON

There is a ring of canals immediately surrounding the axis and
another ring occurs in the cortex. The polyps are retractile
within the verruce. In my specimen all the polyps are retracted.
Spicules are present in the anthocodie.

The spicules of the polyps are (1) small tri-radiate spicules with
sharp spines on the blunt ends of the rays, this being the
commonest type of spicules in the polyps; (2) rods with spines on
the expanded ends, some of which approach a dumbbell shape ;
(3) approximately straight rods; and (4) spindles. The spicules
are arranged in eight longitudinal bands on the anthocodiz and
do not appear to be in any definite arrangement in relation to one
another. They are all red in colour.

The spicules of the axial part of the trunk are (1) colourless rods
with few spines, (2) spindles with blunt processes, the latter with
spines or thorns, and (3) irregularly-shaped spicules.

The spicules of the cortex of the trunk are most frequently
spindles (with a similar shape to those of the axial part), tri-
radiate spicules, short rods, and small irregularly-shaped spicules.
The spindles are more characteristic of the cortex, and the rods of
the axial part. The wall of the stomodeum contains numerous
small red spicules, mostly tri-radiate forms.

The dimensions of the spicules are as follows :—From the polyps :
(1) large spindles, from 0-119 x 0-085 to 0-188 x O19) mamas
(2) tri-radiate forms, from 0:051 x 0:0357 to 0-068 x 0-068 mm. ;
(3) 4rayed forms, from 0:051 x 00425 to 0-068 x 0-051 mm. ;
(4) small dumbbell-like forms, but with double ends, about
0-051 x 0:020 mm. .

From the cortex of the trunk: (1) large spindles, from
0:0935 x 0:0595 to 0:221 x 0-119 mm ; (2) small tri-radiate forms,
from 0:0357 x 0-051 to 0:0595 x 0:022 mm.; (3) dumbbell-lke
forms approximately the same size as those of the polyp.

From the axial part of the trunk: (1) large spindles (only a few
occur), from 02040-1105 to 0:289 x 0:0935 mm.; (2) rods,
from 0°085 x 0°0255 to 0°272x0:0765 mm.; (3) irregularly-
shaped spicules, from 0-238 x 0-119 to 0:340 x 0-119 mm.

Locality, ete.—‘ Pieter Faure,” No. 13139. Cape Morgan,
N.N.E. 92 miles. Depth, 47 fathoms. Procured by dredge.
Nature of bottom, broken shells. Only one specimen of this
species was collected.

Studer gives the following diagnosis of the genus Suberta :—

“Stamm einfach oder verzweigt, aufrecht, mit einer Achse, die
aus unverschmolzenen, von Hornsubstanz umgebenen stabformigen
Spicula gebildet wird und der Ernihrungscanale entbehrt. Rinde
dick, enthalt spindelférmige stachlige Spicula, Die Polypenwarzen
sind gross, senkrecht vom Stamme abstehend, die Offnung an der
Spitze der Warzen achtstrahlig. Die Polypen von der Basis bis
in die Tentakel mit feinen spindelformigen Spicula erfwllt. Um
die Achse ein Kranz von Langscaniilen.”

Studer adds that the genus Suwberia stands near Spongioderma,
but differs from it in the form of the calyx and spicules.

CORALS FROM SOUTH AFRICA.

873

Three species of the genus Suberia have been described, and the
following table shows the differences between them so far as these
can at present be stated.

Suberia capensis, sp. n.

Colour, dark red.

Simple, unbranched.

Height 53 mm.

Trunk cylindrical,
expanded at the apex.

Suberia genthi Str.

Branched.
80 mm.
Cylindrical.

Polyps at the apex but Polyps not terminal.

not terminal, 2-3 mm.
in height.

The rind is firm and
rough.

The spicules are :—
(1) rods, (2) spindles,
(3) tri-radiate forms,
(4) 4-rayed forms,
(5) irregular spicules.

No canals in axis.

(1) straight warty
spindles,
(2) curved warty
spindles,

(3) 4-rayed forms,

(4) double crosses,

(5) irregularly branch-
ed spicules.

Canals in the axis.

Suberia koellikeri Str. Suberia clavaria Str.

Rose-red, disappearing Flesh-coloured during
in alcohol. life, disappearing in
alcohol.

Branched. Simple, unbranched.
About 110 mm. 160 mm.
Cylindrical, branches Cylindrical, expanded
expanded at the at the apex.
apices.

Polyps thickly crowded
together at the apex,
1-2 mm. in height.

The rind is thin and The rind is soft and

1-15 mm. in height.

rough.

(1) rod-like spicules,
(2) cross-shaped or
triplets.

smooth.

(1) slightly spined
spindle-shaped rods,
(2) crosses,

(3) spinous spindles.

Canals immediately out- Canals in the circum- Axis surrounded by a

side the axis.

Axis of horny tissue with
numerous associated
spicules.

Dimensions of Spicules :—
Large spindles, from
0:0935 X 0°0595_ to
0'0289 X 0'0935 mm. ;

tri-radiate forms, from
00357 X 0:01 to 0°068
<0°068 mm.

rods, from 0°085 X 0°0256
to 0°272X0:0765 mm. ;

irregularly - shaped
spicules from @'238X
0119 to 0:340X0119

mm.
dumbbell-shaped forms
about0°051 X 0°020 mm.

Habitat :—South Africa,
Cape Morgan, N.N.E.,
92 miles. Depth, 47
fms. By dredge.
Bottom, broken shells.

ference.

Straight, warty spi-
cules, from 0°09 X
0°08 to 0'34X0°10
mm.

curved, warty spindles
from 0°24X012 to
0°30X0'10 mm. ;

long, spiny, curved
spindles, from 0°'4X
0°04 to 0°24 XK O04
mm.;

irregularly branched
spicules, from 0:24
to 0°06 mm. in widest
diameter ;

4-rayed forms, 0°32 to
0:06 and 0°20 to 0°12
mm, ;

double crosses, from
0'1 to 0°025 and 0°08
to 0°01 mm.

Shallow water, off
Port Jackson
(Australia).

wreath of canals.

Axis consists of smooth Axis as in the last
rod-like spicules species “* well separ-
(with few spines) ated” and consists of
lying close to one long, slightly spined,
another. spindle-shaped rods ;

crosses also occur.

Spinous spindles from
the rind, 0°71 to 0°18
mm. in length ;

Warty, thorny spi-
cules, 0°24 and 0°021
mm. in length ;

rods (‘ Drillinge”)
from axis, 0°35 mm.

long, slightly spined,
spindle-shaped rods,
0°237 and 0°265 mm.
in length.

North of New Zealand. Atlantic, off Monte
Video

874 DR. J. STUART THOMSON ON

SPONGIODERMA VERRUCOSUM Mobius.

This species has already been described from South African
waters by Hickson in his paper on the Alcyonaria and Hydro-
corallinz: of the Cape of Good Hope, Part I.

Localities, etc.— Apparently a shallow-water form.

“Pieter Faure.” 12310. Port Shepstone, north, 8 miles.
By dredge. Depth, 36 fathoms. Nature of bottom, broken shells
and stones. Date, October 30, 1902.

‘“‘ Pieter Faure,” 11127. Umkomass River mouth, N.W. by W.
2 W.54 miles. By large dredge. Depth, 40 fathoms. Nature
of bottom, broken shells and stones. Date, December 30, 1900.

Family MeLITODIDA.

MELITODES ESPERI W.&S8. (Text-fig. 167.)

The specimen is about 104 mm. in length and 117 mm. in
breadth, but these dimensions are only approximate as several of
the terminal branches were broken away. The colony had three
beautiful young specimens of Gorgonocephalus intertwined among
its branches.

Text-fig. 167.

Melitodes esperi and Gorgonocephalus.

The colony is red in colour, and is attached to the substratuin
by a broad, trunk-like base. The branches are mostly in one
plane, although a few are slightly turned inwards in a direction

CORALS FROM SOUTH AFRICA. 875

vertical to the main axis. There is a slight anastomosis of the
branches; the meshes thus produced are very irregular in shape.
Sometimes branches, which may be slightly divided, are seen
growing into the meshes; these offshoots may arise from the
internodes, but this is rare.

The nodes and internodes are very distinctly marked, the former
being dark red and the latter hght red in colour. The internodes
vary in length from 2 mm. near the base to 9 mm. towards the
apex of the colony. The nodes are more or less ring-shaped, but
they have a slight projection in front. The nodes are smaller
towards the apices of the branches than near the bases and are
not so easily distinguished from the internodes. Near the base
the nodes may be 4 mm. in length and 3 mm. in diameter ;
towards the apex they are about 1 mm. in length and diameter.

The axis consists of long needle-like spicules densely compressed
together. The lower part of the axis is not perforated by canals.
On a branch of average size, the diameter of the axis is about
1 mm. The internodes of the axis are pale pink, the nodes
darker in colour.

The base of the colony is a thick trunk about 5 mm. in
diameter, but it soon divides and gives rise to the branches. The
branches are disposed in a more or less fan-shaped manner.
Various organisms, such as Polyzoa, Brachiopods, and Hydroids,
are thickly grouped at the base of the colony.

The coenenchyma covering the axis is about 4 of a millimetre
in thickness: its surface is densely covered by an extremely large
number of spicules of varied size and shape. A rough surface is
produced by the slightly protruding spicules.

The contracted polyps appear as yellow papille on the surface
of the ceenenchyma, and are about °6 mm. in height and 1 mm. in
breadth. On the large, basal branches of the colony, the polyps
are only distributed on three sides, but on the apical branches
they occur on all sides. Polyps are situated on the nodes, and
not simply confined to the internodes. The spicules of the polyps
are yellow, and are arranged in eight areas.

The spicules of the coenenchyma are of the following forms :—

Clubs, with broad leaf-like processes; curved spindles with
narrow processes, some of these spicules are so much curved as
almost to form a semicircle; straight spindles of two forms:
(a) those with small processes, and (6) those with broad, expanded
processes; other spicules are irregular in shape, owing to the
presence of broad processes, which produce a more or less branched
appearance; one of these spicules has, for example, three apical
prongs or processes.

The spicules of the ccenenchyma are all red in colour.

The spicules of the polyps are as follows :—

Straight spindles fairly similar in form to those of the ccenen-
chyma; curved spindles, also fairly similar in shape to those of
the coenenchyma, some of these spindles are bifurcated at the end

876 DR. J, STUART THOMSON ON

in a similar manner to that shown by Wright and Studer for this
species ; straight rod-like spicules with rounded processes ; clubs
and spicules of an irregular shape.

The spicules of the polyps are yellow in colour.

The spicules of theccenenchyma have the following dimensions :—
(a) Clubs, from 0:102 x 0:054 to 0:187 x 0-076 mm. ; (6) curved
spindles, from 0-085 x 0:0204 to 0:238 x 0-047 mm. ; (c) straight
spindles, from 0°0765 x 0:0204 to 0-272 x 0:0646 mm. ; (d) irregu~
larly-shaped spicules, from 0-085 x 0'068 to 0:119 x 0-068 mm.

The dimensions of the polyp spicules are as follows :—(a) straight
spindles, from 0°081 x 0-037 to 0°153x0:025 mm.; (6) curved
spindles, from 0:122 x 0-084 to 0°187 x 0-034 mm. ; (c) clubs, from
0-085 x 0:034 to 0-136 x 0-051 mm.; (d) irregular spicules, from
0:076 x 0:110 to 0°170 x 0:034 mm.

Locality, etc—P.F. 852. About 25 miles east of Hast London.
Tat, 32° 48’ 30" S.; Long. 28° 11! 15” KE. By shrimp trawl.
Depth, 22 fathoms. Nature of bottom,mud. Date, January 11,
1899.

Me.iroprs noposa W. & 8.

The specimen was not complete, the base being absent and many
of the terminal branches had become broken away. ‘The entire
colony probably measured 230 mm. in length and 120 mm. in
breadth. The branches all arise in one plane. The nodes are
very prominent, measuring as much as 6 mm. in diameter and
5 mm. in height near the base of the colony, and 2 mm. in
diameter and 1-5 mm. in height on the small apical branches.
The internodes vary considerably in length, namely, from 2 mm.
near the base of the colony to 12-13 mm. on the terminal
branches. The internodes are more or less cylindrical, and their
diameter varies from about 3 mm. near the base to | mm. at the
apex. Several of the branches originate from the internodes, as
in the specimens of this species collected by the ‘Challenger.’
There is a slight anastomosis near the base of the colony, produced
by branches springing from the nodes.

On the upper branches the polyps are distributed on all sides ;
on the lower part, however, they are mostly confined to three
sides. The polyps are retractile within well-defined verruce,
which measure about 1 mm. in diameter and 0°75 mm. in height.

The ground colour of the colony is yellow, the nodes are brown,
the verruce yellow. The axis, when deprived of the overlying
cenenchyma, is brown at the nodes and white at the internodes.

The conenchyma is rough owing to the presence of numerous
spicules and has a thickness of 1-2 mm. ‘The surface of the
cenenchyma has longitudinal grooves. The spicules of the
cceenenchyma are (1) irregularly-curved spindles, (2) irregularly-
branched spicules, (3) curved, spinous spindles, (4) 4-rayed forms.
‘heir dimensions are as follows :—Irregularly-curved spindles,
from 0:204x0:059 to 0:238x0:051 mm.; irregularly-branched
spicules, from 0'0765 x 0:068 to 0°136x0-085 mm.; curved,

CORALS FROM SOUTH AFRICA. 877

spinous spindles, from 0°153x0-025 to 0170x0°025 mm. ;
4-rayed forms, 0-051 x 0-051 mm.

The spicules of the polyps are (1) curved spindles, (2) straight
spindles, and (3) irregular spicules. The dimensions of these
spicules are as follows:—Curved spindles, from 0:136 x 0017 to
0-250 x 0°034 mm.; straight spindles, from. 0°023 x 0°051 to
0-255 0-056 mm.; irregular spicules, from 0-068 x 0-051 to
0-170 x 0-085 mm. The spicules of the polyps may be contrasted
with those of the ccenenchyma by the predominance of straight
and curved spindles.

The ‘Challenger’ specimens were collected (1) off the New
Hebrides at a depth of 60-120 fathoms, (2) on Hyalonema-ground,
off Japan, at a depth of 345 fathoms. These specimens differ
from mine in the colour of the cenenchyma and axis; the
cenenchyma being reddish brown and the axis yellowish red.
In both, the polyps were yellow and the nodes darker than the
internodes. ‘The shape of the spicules of my specimen rather
differs from that of the ‘Challenger’ forms, but, on the other
hand, the spicules seem to vary so much within this species that
I do not feel inclined to lay great stress on this point.

Locality, etc—P.F. 18381. Off Flesh Point, N. 6 miles to
Flesh Point, N. 2 W. 64 miles. By large trawl. Date,
January 15, 1904.

3
MELITODES DICHOTOMA Pall.

Fragments of a red variety of this species. This species has
already been recorded from the Cape. It is interesting to note
its occurrence in such shallow water, namely from 6—14 fathoms.
At Gordon’s Bay, in all probability it may be thrown up on the
shore during storms, as is so much the case with Gorgonia lammea
HE. & 8. iz

Locality, etc—P.F. 15725. Off Gordon’s Bay. By dredge.
Depth, 6-14 fathoms. Nature of bottom,rock. Date, October 20,
1902.

Family Isip &.

CrRAToISIS RAMOSA Hickson. (Plate XLIII. fig. 1.)

This beautiful form has already been described by Hickson
from South African waters. The species is very fragile, and the
delicate terminal branchlets are very liable to become broken
away. In Hickson’s specimen the base was wanting, but my
specimens are more complete in this respect. The calcareous base
measures about 8 mm. in length, 4 mm. in breadth, and 2 mm. in
height.

The height of the complete specimen would be about 100 mm.
The first part of the main stem immediately above the base is
brown or bronze in colour.

My specimens agree with Hickson’s description of forms col-
lected off Vasco da Gama Peak at a depth of 230 fathoms.

The distribution of colour on the colony appears to vary. The

878 DR. J. STUART THOMSON ON

ivory-white internodes, as described by Hickson, appear to become
reddish in the upper part of the colony, but this is apparently not
always the case.

Locality, ete.—P.F. 13197. Cove Rock, N.W. ? W. 134 miles.
By dredge. Depth, 80-130 fathoms. Nature of bottom, coral
and rvecks. Date, July 30, 1901.

Family MuRICEID&.

Muricerpes FuscA, sp.n. (Plate XLIV. figs. 4 a—c.)

The colonies are about 102 mm. in height and 95 mm. in
breadth. The branches are all in one plane. The main trunk
has a strong horny axis and an expanded base measuring 14 mm.
in length and 12 mm.in breadth. The main trunk has a diameter
of 5mm. and gives off its first branch 13 mm. from the base.
The branches of the first order are given off irregularly from the
trunk, and these, in their turn, give off branches of the second
order, which may also bear small terminal branches. ‘The
branches of any order do not, as a rule, originate opposite one
another. The axis of all parts is thick, the cenenchyma thin.
The axis is brown near the base, and light yellow at the apices.

The main trunk is fairly cylindrical, but the branches are
slightly flattened. This flattening of the branches appears to be
due to the manner in which the ccenenchyma grows over the
axis, as the latter is fairly cylindrical in shape. The upper branches
are thin and flexible, but their apices are expanded. The polyps
are mainly confined to three sides of the axis. The general
surface of the ccnenchyma is covered by small, protruding
spicules. These spicules do not overlap, but there are only
minute spaces between them.

The polyps are also well protected by spicules. There is a
crown of spindles, in which the spicules are disposed en chevron,
in eight triangular areas which are very pointed towards the
apices. In each of these triangular areas there appear to be
about 8-12 spindles, which are placed more or less vertically to
the surface. At the base of those areas the spindles are arranged
in a continuous ring surrounding the polyp. The spindles
forming the ring also overlap one another, but not to such an
extent as those in the triangular areas. There appear to be about
ten spicules at any one place in this ring, enumerating in a
proximo-distal direction. Distally to this ring, spicules of
another form are situated on the polyp wall. These are very
protuberant, and stand more or less in a vertical direction.
These spicules are ‘“* Blattkeulen” and pass over basally into small
spicules of the same general form as those of the general surface
of the ccenenchyma, namely “‘ Kalkkorper.” The entire polyp is
thus well protected by spicules.

The polyps are capable of being withdrawn within the verruce,
but the latter are small and do not rise any distance beyond the
surface of the cenenchyma. The polyps vary considerably as to

CORALS FROM SOUTH AFRICA. 879

the degree of extension, but the apices of the tentacles are never
seen to any extent.

In all the better expanded polyps, the crown of spindles is very
apparent. This crown has frequently a reddish or brownish
coloration. The general surface of the colony has a greyish
colour, which is produced by the spicules.

Owing to the varying degree of extension, the polyps naturally
vary considerably in size,

The verruce have minute lobes, measuring as much as 2 mm.
in height and 1:6 mm. in diameter.

The ceenenchyma is thin at all parts and sometimes is 1 mm. in
thickness.

The axis of the secondary branches varies in diameter, from
about 1:3 mm. in the lower branches to 1 mm. in the apical
branches.

The interval between the polyps varies from about -1 mm.
to 19 mm. There is a terminal polyp at the apices of the
branches.

The spicules of the crown of the polyp are straight or curved
spindles, armed with blunt tubercles. The majority of the
spindles are very much curved, but a few are straight. The
tubercles have a tendency to great irregularity of shape, and
vary very considerably. The spicules from this part also vary
notably in shape, some being rod-like, others club-like, but
there are all transitions between the different forms. The
number of tubercles or spines also varies considerably, in some
spicules there are scarcely any, in others they are numerous. In
some of the more curved spindles there is a secondary offshoot
from the main axis about the middle of its length.

The spicules of the lower part of the polyp differ, as a rule, in
shape from those of the upper crown. Many of the spicules from
this part are “DBlattkeulen,” others are more of the form termed by
Kolliker “ Kalkkorper.’ There is much variety in the shape of
those spicules, probably every stage grading the one into the
other. There are also a few spicules similar to those occurring in
the crown of the polyp, namely curved spindles. There are other
spicules which may be more appropriately termed spinous clubs.

The superficial spicules of the coenenchyma are much smaller
than those of the polyp. They are mostly small “ Aalkkorper”
with irregular blunt processes coming off in all directions, less
frequently there are short spindles with broad processes.

The dimensions of the spicules are as follows :—Spindles of the
polyp crown, from 0:102 x 0-017 to 0°340 x 0:085 mm.; ‘ Biatt-
keulen” of the lower polyp, from 0°17 x 0°119 to 0-289 x 0°085 mm. ;
“ Kalkkorper” of the lower polyp, from 0°153 x 0-119 to 0°204 x
0:153 mm.; ‘“ Kalkkorper” of the eenenchyma, from 0-047 x 0-030
to 0105 x 0:076 mm. ; spindles of the cenenchyma, from 0:030 x
0:023 to 0-088 x0-061 mm.

There was unfortunately no bel attached to this specimen,
and thus the exact locality cannot be stated.

880 DR. J. STUART THOMSON ON

Nutting has given the following diagnosis of the genus
Muriceides :—

“ Muricemes W. & 8. (emended).

Muriceides Studer + Clematissa Studer, Archiv f. Naturgesch.,
Jahrg. liii. Bd. i. pp. 54, 55. .

Muriceides Wright & Studer + Clematissa Wright & Studer,
‘Challenger’ Reports, the Alcyonaria, 1889, pp. lu, lin, 105,
106.

‘“‘Calyces cylindrical, or in the form of truncated cones, their
walls filled with vertically-placed spindles, often modified into
clubs, discs, or triradiate forms vertically placed, but not ‘en
chevron,’ and not forming a true crown of points around the
margin. The opercular spindles are placed ‘en chevron’ on the
tentacle bases. The ccenenchyma contains spicules of various
forms, and the branches may, or may not, end in calyces.”

ACANTHOGORGIA ARMATA Verrill.

The colony is irregularly branched. The ccenenchyma is thin,
and filled with conspicuous, white spicules. The verruce are
elongate, often curved, capitate or clavate, surmounted by eight
groups of long, divergent, sharp spicules, with an irregular
““chevroned ” arrangement.

The axis has a yellowish-brown colour. The spicules are white,
rough, curved, and fusiform. The colour of the colony is ash-

rey.

; This species has been previously recorded by Hickson from
off Vasco da Gama Peak, 230 fathoms. There are three good
examples in the present collection.

Locality, etc—P.F. 18857. Cape Morgan, N. 2 W. 13 miles.
Depth, 250-320 fathoms. By shrimp trawl. Nature of bottom,
broken shells. Date, July 8, 1906.

Family PLEXAURID2.

EUNICELLA PAPILLOSA Esper. (Plate XLIII. figs. 2 & 3.)

The colony measures 162 mm. in length and 112 mm. in
breadth. It has a basal encrusting part measuring 11 mm. in
length and 15 mm. in breadth. The branches are all disposed
in one plane. The verruce occur over the entire surface of the
colony, even on the flat encrusting base. The main axis and
most of the branches are cylindrical, but occasionally some of the
upper branches are slightly flattened. Proceeding from the main
stem, the branches gradually decrease in diameter but expand
again near the apices. Some of the branches have a few local en-
largements, but these are probably not of specific importance.
The verruce give to the colony its papillated appearance; the
largest are about 2 mm. in height and 1:2 mm. in diameter,
the smallest are only fractions of a millimetre. The surfaces of
the verruce are covered bya mass of spicules. A rough transverse

CORALS FROM SOUTH AFRICA. 881

section through a branch shows (1) the horny brown axis in the
centre, (2) a ring of canals grouped round the axis, and (3) ex-
ternally, the cavities of the polyps arranged in a circumferential
series.

The spicules are situated on the verrucz, on the septa between
the polyp cavities and surrounding the horny axis. These spicules
are very abundant and are as a rule placed in a radial direction ;
they are of two kinds, viz. (1) the characteristic torch-like spicules
and (2) spindles. The torch-like spicules measure about 0-068 x
0:025 and the spindles from 0°0935 x 0:034 to 0-136 x 0°0374 mm.

A portion of a colony of Hunicella papillosa was dredged in
Simon’s Bay during the voyage of the‘ Challenger,’ and Hickson
has also recorded it from Rij Bank (Algoa Bay) at a depth of
25 fathoms. Hickson also draws attention to the remarkable
resemblance between the spicules of Gorgonia albicans and those
of Hunicella papillosa, more especially in regard to the torch-like
spicules.

He says, “The examination of my preparations of spicules
alone would lead any one to the conclusion that they were taken
from the same species. Yet the specimen of Hunicella papillosa
obtained on Rij Bank, 25 fathoms, has a cylindrical axis and
prominent verruce, whilst the specimen of Gorgonia albicans
picked up on the beach at Port Alfred has a very much flattened
axis, thin cenenchyma, and inconspicuous verruce.”

Hickson thinks that there is justification in transferring
Eunicella papillosa to the family Gorgoniide,and that Lunicella
papillosa and Gorgonia albicans should probably be included in
the same genus.

Locality, ete.—P.F. 15801. Off Seal Island, 8.S.E. 24 miles,
and Seal Island, S. by E. 2} miles. Procured by dredge. Depth,
9-10 fathoms. Nature of bottom, broken shells. Date, October 30,
1902.

PSAMMOGORGIA PULCHRA, sp.n. (PI. XLIII. fig. 5; Pl. XLV.
figs. 3a & b.)

This very beautiful form has an orange ground-colour, with
the red tips of the tentacles projecting. The colonies are not
all complete. A small colony (P.F. 858) consists of a fairly
flat, expanded, basal part which gives rise to two upright shoots,
one of which branches into two, the other being simple. The
basal part of the colony is 11 mm. in length, 6 mm. in breadth,
and 3mm. in height. The simple, upright shoot is 9mm. in
height, 2 mm. basally and 41mm. apically in diameter. The other
(divided) shoot is 28mm. in height, 3mm. in diameter at the
base and 5mm. at the apex. The branches are thus expanded at
the apex. Another specimen (15345), which is probably not quite
complete, has a very small basal part. The base gives rise to a
main shoot (2mm.in diameter), which at a distance of 9 mm. from
its origin sends off a lateral branch about 12 mm. in height; a
second small branch is given off from the main shoot after an

882 DR. J. STUART THOMSON ON

interval of 12 mm. This second branch measures 11 mm. in
length and 3 mm. in diameter.

The main shoot is 21 mm. in height and has a diameter of
5—6 mm. at the apex.

The surface of the cenenchyma is covered with large, yellow
spicules, which are easily seen with the naked eye. They lie ad-
jacent to one another but do not overlap, the boundaries of each
being clearly defined. These superficial spicules of the ccenen-
chyma do not appear to be arranged in any very definite manner
in relation to one another; they are of such varied sizes that any
great regularity is prevented. It may, however, be observed that
all the superficial spicules of the coenenchyma have their long axes
in the direction of the longitudinal axis of the colony; in the
verruce, the spicules have a corresponding position in relation to
the polyps.

The spicules of the cortex are (1) broad spindles, yellow in
colour, (2) long narrow spindles, of a pale colour. The spicules
project very much on the surface at all parts. The spicules of
the anthocodie are red and are long spindles arranged ‘ en chev-
ron’ in eight longitudinal, triangularareas. At the base of these
strips, a layer of spicules forms a ring round the polyp. This ring
is composed of two or three spicules lying one behind the other in
a proximo-distal direction.

The verruce have a circle of projecting spicules, only one layer
deep. This circle consists of 14-16 yellow spicules which lie with
their long axes parallel to the length of the polyps. These pro-
jecting spicules appear to be more prominent towards the apex of
the colony. The general appearance of the polyp spicules reminds
one, at first sight, of the setting of a stone in a finger-ring; the
anthocodia with its red spicules resembling a jewel which is
surrounded by little rivets, namely, the large, yellow spicules of
the verruce. The degree to which the anthocodize are exposed
beyond the verruce varies to some extent; as a rule, eight trian-
gular areas are easily seen, in other cases there is only a minute
area, with spicules or only an opening at the apex of the verruce.
The axis is horny in the centre, but with long narrow needles or
spindles on its surface. ‘These axial spicules are about 0-340 mm.
in length and 0:051 in breadth ; their margins are almost smooth.
Tn a rough transverse section, the axis is seen to be composed of
a large central grey area surrounded by a brown ring. The axis
is not penetrated by canals.

The polyps occur on all parts of the colony including the basal
encrusting part; they are arranged in a spiral manner on the
branches. The polyps are about 15mm. in height and 2 mm. in
diameter.

The dimensions of the spicules are :—(1) large, yellow spindles,
from 0-081 x 0:039 to 0-123 x 0-021 mm.; (2) small, yellow
spindles, from 0-042 x 0-024 to 0-093 x 0°033 mm.; (3) red spindles,
from 0:021 x0:0015 to 0°114x 00135 mm.; (4) long needle-like
spicules of the axis, 0°340 mm. in length by 0-051 mm. in breadth.

CORALS FROM SOUTH AFRICA. 883

Localities, etc.—P.F. 13345. Off Cape Morgan, N.N.W. 7
miles. Depth, 52 fathoms. By dredge. Nature of bottom, rocks,
sand, and shells. Date, August 12, 1901.

P.F. 858. Off and east of Cape Morgan. Depth, 36 fathoms.
By dredge. Nature of bottom, stones. Date, January 12, 1899.

Verrill’s diagnosis of the genus Psammogorgia is as follows :—
*Corallum dichotomous or subpinnate, with round branches.
Axis horn-like. Coenenchyma moderately thick, the surface finely
granulated with small rough spicula. Cells scattered, sometimes
flat, more frequently raised in the form of rounded verruce.
Polyps with rather large, elongated, slender warty spindles at the
bases of the tentacles. Spicula of the ceenenchyma mostly short,
thick, and very rough, warty spindles and rough, warty clubs of
moderate size.”

Eleven other species of Psammogorgia have been described.

KUPLEXAURA MEDIA, sp. n. (Plate XLIV. figs. 2 a-c.)

The colour of the colony is bright red, with white polyps. A
large part of the colony is concealed beneath a thick growth of
Hydroids, and thus the exact dimensions of the specimen are
rather uncertain. It was probably about 25 cm. in length.

In the lower part of the colony the axis is much flattened, but
towards the apex it becomes rounded. The large axis near the
base of the colony is 3°5 mm. in one diameter and 1:6 mm. in the
narrow diameter, The axis is very small towards the apex of the
colony, and is surrounded by the canals in a circular manner.
The axis is hollow, and has the structure described by Wright and
Studer for the genus Huplexaura.

The base of the colony is missing. The branches are all in one
plane and vary in diameter at different parts of their length.
They are thick near the base, narrower medianly, and expand
again near the apex.

The polyps are arranged in an irregular spiral round the
branches. The polyps are in many cases fairly well extended, but
others are retracted and appear only as minute pores on the
surface of the cenenchyma. These pores vary in shape, some are
circular or oval, others are slit-like and elongated in the direction
of the long axis of the branch. The pores are from °3 to ‘5 mm.
in diameter. A well-expanded polyp is about 1-2 mm. in length
and *5 mm. in diameter.

Each tentacle has about ten pairs of pinnules. The spicules are
situated at the bases of the tentacles and extend in a distal
direction. There is, firstly, a series of spicules on the polyp sur-
face in a line with the tentacles, and, secondly, another series
placed intermediate to the first. The second series has not so
many spicules as the first. The spicules in each of these series are
not arranged in a very regular manner.

The cenenchyma is dense and granular. The spicules are
arranged in a very dense layer in the cortex, but there are fewer
towards the axis.

884 DR. J. STUART THOMSON ON

The spicules of the cortex are mostly spindles with irregular,
blunt processes. Some have a simpler form than others. The
spicules of the centre resemble those of the cortex, but simpler
forms occur such as rods (with small processes) and crosses. ‘The
spicules of the polyps are very minute, needle-like or rod-like
spicules, with blunt processes.

The dimensions of the spicules are as follows :—Spindles of the
cortex, from 0:059 x 0-034 to 0°085 x 034 mm.; spindles of the
centre, from 0:051 x 0:034 to 0:085 x 0°051 mm.; rod-like spicules
of the polyps, from 0-064 x 0-008 to 0:112 x 0-034 mm.

This species resembles Hupleaaura brauert Kiikenthal, /. albida
Kiikenthal, and Z#. parciclados Wright & Studer. It has the
closest resemblance to Huplexaura brauert Kiikenthal, the spicules
of which are almost identical with it except in size. My specimen
differs, however, from Huplexaura brauert in the arrangement of
the polyp spicules and in other points. From the shape of the
spicules alone, one would probably conclude that they belonged to
the same species.

Locality, etc.—P.¥. 742. Between Roman Rock and Cape
Recife. Depth, 17 fathoms. By dredge. Nature of bottom,
corals. Date, December 12, 1898.

Family MALACOGORGIIDS.

MALACOGORGIA CAPENSIS Hickson.

The colony consists of a main stem (with an expanded, basal,
attaching part) which divides at some distance from the base into
two primary branches; one of the latter gives rise to three, the
other to eleven secondary branches. ‘The polyps are situated on
the secondary branches. ‘The main stem is cylindrical and
measures 50 mm. in length by 4mm. in diameter. The basal,
attaching part has a diameter of 6 mm. The polyp-bearing
branches are long and slender, the longest in my specimens being
80 mm., the shortest 27 mm. in length.

The branches originate in the manner shown in Hickson’s
figure, coming off right and left in one plane.

There are no polyps at the bases of the secondary branches, thus
leaving bare areas, 3-7 mm.in length. The younger polyps are
situated near the bases of the branches. The polyps are fully ex-
panded ; a large one measures 1 mm. in length and about °75 mm.
in diameter. The general appearance of the polyp reminds one
superficially of a contracted Hydra. On the lower part of the
branches the polyps have a bilateral arrangement, but higher up,
and especially near the apices, they originate on all sides and form
a dense cluster.

The tentacles are about 0:4 mm. in length and have 12 pairs of
pinnules.

The axial part of the secondary branches has a diameter of 0:5
to |] mm.

CORALS FROM SOUTH AFRICA. 885

IT am able to confirin Hickson’s statement regarding the absence
of spicules in this genus. My specimens are larger than that
described by Hickson, and the polyps in his form are also smaller.
My specimens, however, agree well with his description.

Hickson gave the following diagnosis of the genus J/alaco-
gorgia :—

“Colony slightly branched. Axis horny with no trace of lime.
No spicules in any part of the colony. Pelyps arranged bilaterally
in the plane of branching at the basal two-thirds of the secondary
branches and on ali sides of the terminal one-third of the
secondary branches.

“ Malacogorgia capensis, with the characters ef the genus.
Colour in spirit, white.”

Hickson placed this form in a new family, the Malacogorgiide.
He gave the characters of the new family as follows :—‘“ Colony
branched and upright. Axis slender, horny. Spicules and all
other forms of calcareous skeleton absent.”

Localities, etc. P.F. 18729. Bird Island (near Cape Seal),
EK. by N. + N. 5 miles. By large trawl. Depth, 40 fathoms.
Nature of bottem, mud. Date, August 29, 1905.

P.F. 703. Lat. 33° 53! 15" S., Long. 25° 51! 45" EK. By large
trawl. Depth, 26 fathoms. Nature ef bottem, mud.

Family Primnorps#,

STACHYODES GILCHRISTI, sp.n. (PI. XLIV. fig. 1; Pl. XLV.
figs. 2a & 6.)

The specimens are not complete. The branching is in one plane,
but is neither dichotomous nor regular. The polyps are arranged
in verticils which-are separated from one another by an interval
of about a millimetre, but the degree ef separation varies con-
siderably at different parts ef the celony. Thereare onan average
five verticils te every ten millimetres. The length of a large
verticil is about 3 mm. on the ad-axial and 2°5 mm. on the ab-
axial side. The verticils are about 5°5 mm. in diameter,

The axis, which is brown on the lower part of the colony and
yellow nearer the apex, is covered by sclerites which vary con-
siderably in size and shape. These sclerites covering the horny
axis are thin and fairly trausparent. The diameter ef the axis
is about *35 mm.

In each verticil there are usually five polyps. The polyps are
protected by three pairs of sclerites, namely an ad-axial pair,
amedian pair, and an ab-axial pair. The ad-axial pair consists of
long sclerites which are produced outwards in a lateral direction,
the median pair are smaller. The ad-axial and median spicules
have sharp, rather angular margins, and when the latter are
viewed from the side, they appear like spines. The ab-axial pair
of sclerites have slightly ribbed or dentate margins. The ad-axial
sclerites of the two lateral polyps form a ring or tunnel embracing

Proc. Zoou, Soc.—1911, No. LXI. 61

886 DR. J. STUART THOMSON ON

the main axis. The sclerites vary considerably in size, but they
probably range from 0:085 x 0:153 to 4:0 x 2:0 mm.

The diameter of a branch in the intervals between the
verticils is about 3 mm.

The sclerites covering the axis have the following forms :—(1)
triangular, (2) quadrangular, (3) forms with curved sides, and
(4) irregular forms.

The systematic position of this species is near Stachyodes
trilepis Pourtalés and Calyptrophora josephine Lindstrom.

Locality, etc—P.F. 11966. Cape Vidal (Natal), N.N.E. 4 N.
94 miles. Depth, 80-100 fathoms. By dredge. Nature of bottom,
rocks. Date, February 27, 1901.

THOUARELLA HICKSONI, sp. n. (Pl. XLIV. figs. 3a & 5;
TP; ROU die, Js)

The colony is more or less bottle-brush-like, but it gradually
tapers off towards the apex. Thedimensions of the colonies vary,
the largest specimen was 66 mm. in length and 25-28 mm. in
diameter. The central stem gives rise to the polyp-bearing
branches in a spiral manner.

These branches vary in length, they are longer near the base
of the main stem, and gradually decrease in size towards the apex.
The branches are as a rule simple, but in some cases there are
secondary and tertiary branches. The: primary branches are not
arranged in a regular spiral as the distance between their points
of origin is not the same in all cases. The interval between the
origin of two primary branches on the central stem is frequently
about 1 mm.

The axis is horny, flexible, yellow and iridescent, and its surface
is covered with imbricating spicules. It is more or less oval in
transverse section. When the spicules are removed from the
surface of the axis, fine longitudinal lines or strieareseen. There
are generally three spicules in a transverse row on the axis of a
branch. The diameter of a branch is about 0°391 mm.; that of
the horny axis about 0°136 mm.

Most of the polyps are situated on the primary, secondary and
tertiary branches, but sometimes they occur on the central stem.
On the branches they originate singly, and are disposed in a spiral
manner. ‘The polyps are pear-shaped, and their surfaces are
covered by imbricating scales. The size of the polyps varies con-
siderably, the following dimensions may be noted :—(1) 1-088 x
0°629 mm.; (2) 0-476 X0°306 mm.; (3) 1°343 x 0-595 mm. ; (4)
0:595 x 0°374 mm. ; (5) 0°935 x 0°663 mm. ; (6) 0°612 x 0-391 mm.

The interval on the branches between the polyps also varies but
not to any extent. ‘The distance is generally from °5 to °6 of a
millimetre. In some polyps, the tentacles were extended beyond
the verruce, and the pinnules were seen, though not sufticiently
well to determine their number. There are on an average about
twenty-six spicules on each verruca and these are disposed in six
transyerse (excluding the opercular spicules) and four or five

CORALS FROM SOUTH AFRICA. 887

longitudinal rows. The spicules of the verruce show considerable
variation; they may be grouped in two sets, namely (1) those
covering the lower or general surface of the vérruéz, and (2) the
apical or opercular spicules. The lower spicules of ‘the verruce
are triangular, rectangular, scale-like or plate-like spicules. In
their imbricate ar rangement and also in their shape, these spicules
remind one of the scales of fishes. Their surface is marked, except
near the margin, by a number of small, rounded pores, which are

arranged more or less in rows running in the direction of the long
axis of the polyp. The margins oe these scale-like spicules is
frequently, wholly or partially dentate. The opercular spicules
situated at the apices of the verruce differ in shape from those
last described. They are more or less triangular, with a
pr ojecting spine in front, the entire spicule resembling a Skate
(Raia) inform. These oper cular spicules are éight i in number and
their spines project towards the central opening of the verruca.

The surface of these opercular spicules have pores similar to those
of the lower part of the verruca.

The dimensions of the spicules are as follows: a) lower
spicules of the verruce from 0°1105 «x 0°1615 to 0°187 x 0:221 mm.
(>) the apical or opercular spicules (including the spine) nee
0-289 x 0°136 to 0°561 = 0°255 mm.

The spiculés covering the main stem of the colony are similar to
those of the lower part of the verruce, but are frequently much
smaller. They have a more or less ir regular arrangement and do
not as a rule everlap oné another. Their size is very variable,
ranging from 0-068 x 0-025 to 0:221 x 0°153 mm.

Around the basal part of the colony a Polyzoan is encrusted,
and intertwined among the upper branches are small Ophiuroids.

Locality, éte.—P.¥. 14265. Off Cape St. Francis, N.E. by E.
32 miles. By dredge. Depth, 74 fathoms. Nature of bottom,
rocks, Date, February 19 19, 1902.

Family GoRGONTID A

GorGoniIA CAPENSIS Hickson.

This is a béautiful example of a species described by Hickson
from the Cape in 1900.

Hickson’s specimen is larger, namely 250 mm. in length, while
my example is only 140 mm. long.

The spicules are warty spindles (*“ Doppelspindeln” of Kolliker).
The ENTE length of these spindles in Hickson’s specimen is
0-1 mm. ; in mine they are smaller, namely, from 0-0544 x 0:034
to 0° 0935 x 0-034 mm. .

In 1900 Hickson recorded this species as a viviparous Alcyo-
narian, and in 1905 Thomson and Henderson corroborated this
discovery i in specimens from Ceylon.

The Ceylon specimens were collected in deep as well as in
shallow water. They were larger than either of the Cape
specimens, and were practically white. Hickson’s specimen was

61%

888 DR. J. STUART THOMSON ON

collected off Cape St. Blaize, S.W. 3 W. 10 miles, at a depth of
40 fathoms. My specimen was collected farther to the north :—
Between Knysna Head and Nutze River, 3 to 4 miles off shore.
Depth, 35-49 fathoms. By large trawl. Nature of bottom, sand
and mud, The specimen had entertwined among its branches,
the string-like projections of a ‘“‘Mermaid’s Purse” and two or
three Brittle-stars.

GORGONIA FLAMMEA E. & 8S.

This species is collected in great abundance on the shore,
especially after storms, at many parts of the coast. It is
extremely common for example at Gordon’s Bay, False Bay.

Family GORGONELLID&.

ScIRPEARIA FURCATA Hickson, emend. Simpson.

1903. Scirpearia furcata Hickson.
1903. Scirpearia furcata var. ? Hickson.
1903. Scirpearella indica Hickson.
1905. Scirpearia sp., Thomson & Henderson.
1905. Scirpearella sp. B, Thomson & Henderson.
1905. Juncella elongata (Val.) Hickson.
1909. Scirpearella aurantiaca Thomson & Russell.
This species has recently been thoroughly revised by Simpson
and I therefore include his list of synonyms. In preparing his
valuable monograph on the Juncellid Group of the Gorgonellide,
Simpson had an opportunity of examining my specimens from
South Africa.

The South African localities are as follows :—

P.F. 13081. Hood Point, N. 53 miles. Depth, 42 fathoms,
Nature of bottom, saad and shells.

P.F. 858. Offand east of Cape Morgan. Depth, 36 fathoms.
Nature of bottom, stones.

P.¥.12377. Umblangakulu River mouth, N.W. by N.
74 miles. Depth, 50 fathoms. Nature of bottom, sand,
shells and sponge fragments.

P.F. 10841. Umbloti River mouth, N. by W. 2 W. 83 mules:
Depth, 40 fathoms. Nature of bottom, sand, shells, and
hard ground.

PIE 13030: Beacon east of East London, N. 7 E. 10 miles.
Depth, 52 fathoms. Nature of bottom, sand and shells.
P.F. 12033. Cone Point, N.W. 3 W. 4 miles. Depth, 34

fathoms. Nature of bottom, br folkea shells.

P.F. 11543. Tugela River mouth, ING Wey eee

miles.
Depth, 47 fathoms. Nature of bottom, broken shel

2:
]

am Ce)

SCIRPEARTIA FLAGELLUM Studer, emend. Simpson,

1863. Juncella flagellum Johnson.
1864. Juncella flagellum Johnson.
1870. Viminella flagellum Gray.

CORALS FROM SOUTH AFRICA, 889

1881. Scirpearia flagellum Studer

1891. Scirpearia ochracea Studer.

1901. Scirpearia flagellum Studer.

1901. Scirpearia ochracea Studer.

1909. Scirpearia flagellum Thomson & Russell.
T have quoted the preceding synonyms from Simpson’s paper.
The following South African localities at which this species

occurs, may be noted ; ——

P.F. 12855. Buffalo River, East London, N. 15 miles. Depth,
310 fathoms. Nature of bottom, coral and mud.

P.E. 12061, O'Neil Peak, NW 2 W. 92 miles. Depth, 90
fathoms. Nature of bottom, broken shells.

P.F. 12107. O’Neil Peak, N.N.W. + W. 8 miles. Depth, 55
fathoms. Nature of bottom, broken shells.

P.F. 11586. Amatikulu River mouth, N.W. 2? N. 20 miles.
Depth, 62 fathoms. Nature of bottom, rocks and sponges
(hard ground).

HICKSONELLA SPIRALIS Simpson.
Juncella spiralis Hickson,

This species was described from the Cape by Hickson, later
revised by Simpson and named /icksonella spiralis. Unfortunately
the generic name, Hicksonella, is already preoccupied for an
entirely different Aleyonarian genus.

Simpson has given the following diagnosis of this species :—
“Colony unbranched; in the larger forms spirally twisted. The
ecenenchyma is thin and densely packed with scale-like spicules ;
the axis is composed of concentric laminz of a horny substance
in which a calcareous deposit is embedded. The polyps are re-
stricted to a region occupying two-thirds to three-fourths of the
circumference of the ceenenchyma,; a longitudinal bare tract oc-
cupies the remaining part. The verruce are long and club-shaped,
and are evidently not retractile into the coenenchyma; they are
closely packed together, and are covered with minute overlapping,
seale-like spicules. The flat thin scales on the aboral surface of
the tentacles form a sort of pseudo-operculum to the partially
retracted polyp.

The chief types of spicules are :—(1) In the ceenenchyma, very
thick spindles with close-set irregular warts, passing by gradual
transitions to almost spherical warty forms: (2) in the polyps, (a)
long thick spindles with a few long warts, (}) irregular forms and
crosses, (c) small, flat, thin scales.”

Localities, ete —P.F. 13152 4. Cape Morgan, N. 3 W. 104
miles. Depth, 77 fathoms. Nature of bottom, rocks aid broken
shells.

P.F. 13121. Cape Morgan, N.N.E. 92 miles. Depth, 47
fathoms. Nature of bottom, broken shells.

Hickson’s specimens were also collected off Cape Morgan, at a
depth of 36 fathoms.

890 DR. J. STUART THOMSON ON

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Ktxenruar, W., und Gorzawsky, H.—‘ Japanische Gorgoniden,
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CORALS FROM SOUTH AFRICA. 891

May, W.— Aleyonarien Hamburger Magalhaensische Sammel-
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892 ON CORALS FROM SOUTH AFRICA.

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EXPLANATION OF THE PLATES,

PruatE XLIII.

. Ceratoisis ramosa Hickson. X 1}.

. Hunicella papillosa Wsper. Nat. size.

. Eunicella papillosa, part of branch of (with cephalopod egg-capsule
encircling the branch). X 6.

. Suberia capensis, sp.n. Nat. size.

. Psammogorgia pulchra, sp.n. X 1.

n> Go to a

IAS), eH PIL OIL AVAL,

ae PTT
Zs

9 : Huth lith.eb tap .

SCUENENS) (Ol {ESANOIDIO INGE IE0e 2 NES).

ON THE SKULLS OF CYNODONT REPTILES. 893

Prare XLIV.

Fig. 1. Stachyodes gilchristi, sp. n., part of branch of. X 8.
2a. Hupleraura media, sp. u. Spicules of central trunk.

2b. 33 6 i Spicules of external trunk.
2c. y x Spicules of polyps.
3a. Thouar ella hicksoni, sp. n., verruca of. X 45.
3b. 5 apex of verruca of. x 100.

Aa. Muriceides fusca, sp. n. Spicnles of upper part of polyp.
i 5 +5 Spicules of lower part of polyp.
4e. 5 on - Spicules of coenenchyma.

Pratt XLV.

Fig. 1. Thouarella hicksoni, sp.n. Spicules of verruea.
2a & b. Stachyodes gilchristi, sp. u., sclerites of.
3a. EA hg pulchra, sp. n., red spicules of.

35. 53 » yellow spicules of.
4a. Suberia capensis, sp. n. Spicules of central trunk.
4b. 5 fi 5 Spicules of external trunk.
Ac. oe ne 3 Spicules of polyp.

38. On the Structure of the Skullin Cynodont Reptiles *.
By R. Broom, M.D., D.Sc., C.M.Z.S.
[Received April 10, 1911: Read May 28, 1911. ]

(Plate XLVI.7 and Text-figures 168-180.)

Historical and Introductory.

In 1853 the British Museum received from Andrew Bain the
first known skulls of fossil reptiles with a mammal-like arrange-
ment of the teeth. These ultimately became the types of
Lycosaurus tigrinus and Cynodraco serridens. In 1858 Sir
George Grey presented the skulls which were shortly afterwards
described by Owen as Galesaurus planiceps and Cynochampsa
laniaria. The Galesawrus skull though crushed was nearly
complete, and being so very vemarkably mammal-like Owen
almost immediately described it 1n a paper read before the
Geological Society on 20th April, 1859.

Although for seventeen years nothing further descriptive of
any of the reptiles with a mammal-like dentition was published,
it is necessary to briefly consider some of Owen’s other work in
the interval to clear up a certain confusion of nomenclature. In
1859 Owen gave to the world his famous classification of the
fossil reptiles, and though he formed the Order dAnomodontia
for the South African reptiles of the Dicynodont type, he care-
fully omitted all reference to those reptiles, like Galesawrus and
Cynochampsa, with a mammal-like dentition. When in 1861 he
published his ‘ Paleontology,’ feeling compelled to put the
remarkable Galesawrus somewhere, he made it the type of a
“family ” of the Anomodontia, calling it the Cynodontia, doubtless

* On p. 902 Dr. Broom names a new species, viz. Cynoguathus seeleyi.— Kh pitToR.
+ For explanation of the Plate see p. 925.

894 DR. R. BROOM ON THE

recognising that Galeswurus was in some way related to Dicyno-
don, but hesitating to make a new Order on the evidence of a
single skull. As he still defined Anomodontia as reptiles with
‘ teeth wanting or limited to a single maxillary pair,” it is mani-
fest he did not regard Galesaurus as really an Anomodont.

In 1876, when Owen issued his‘ Catalogue of the South African
Fossil Reptiles ’ he put all the forms with a mammal-hke dentition
into a new Order, the Theriodontia. In 1903 I showed that Owen's
Theriodontia is not a natural order, for it included two groups
which, though agreeing in having the dentition specialised into
incisors, canines, and molars, and possibly the one being ancestral
to the other, were yet so dissimilar that they could not be well
kept together. The more primitive group, which occurs only
in Permian beds, has simple molars, an open Rhynchocephaloid
palate, a transpalatine bone, large angular and surangular bones,
a single occipital condyle, no acromion process, and apparently a
digital formula of 2, 3, 4,5, 3. The higher group, which is known
only from Upper Triassic beds, has usually specialised molars, a
secondary palate asin Mammals, no transpalatine, small angular
and surangular bones, two occipital condyles, an acromion process,
and a digital formula 2, 3, 3, 3, 3. As Cynodontia was the name
first applied to animals of the Galesaurus type, this title should
be retained for the higher group. For the lower forms I proposed
the name 'Therocephalia. The name Theriodontia should be
dropped, as only likely to lead to confusion.

Among the new forms described by Owen in his Catalogue is a
badly weathered small Cynodont skull somewhat resembling that
of Galesaurus and named WVythosaurus larvatus. In 1887 he
described another small but well-preserved skull which he believed
to be an additional specimen of Galesaurus.

Most of our knowledge of the Cynodonts, however, is due to
Seeley, who, as the result of his expedition to South Africa, not
only came across the skulls of many new types, principally in the
collections of Dr. Kannemeyer, Mr. A. Brown, and the Albany
Museum, but for the first time obtained most of the skeleton of
some Cynodonts. In one paper issued in 1896 he described a
very fine skull with most of the vertebral column, the lmb-
girdles, and portions of the limbs of a large carnivorous type,
which he called Cynognathus crateronotus, also a fine skull of an
allied form called Cynognathus platyceps from the Albany Museum
collection. In other papers he described new types of Cynodont
reptiles with flat-topped molars. Of these the best known types
are Gomphognathus, Diademodon, and T'rirachodon. These were
regarded by Seeley as belonging to a distinct Order, which he
called Gomphodontia; but as, apart from the specialisation of
the molars, there are no characters of any importance to distin-
guish the Gomphodonts from the Cynodonts, it seems to me
impossible to regard them as forming more than a Family of
the Cynodontia.

Within the last eight years I have been so fortunateas to come

SKULLS OF CYNODONT REPTILES. 895

across a considerable number of new Cynodonts, mostly collected
by Mr. A. Brown, and also to add a good many facts to our
knowledge of the anatomical structure. Probably the most
important of the recent finds has been the discovery by myself
of the nearly perfect skull which I have called Bauria cynops.

Of all extinct reptilian groups there is probably no one of
greater interest than the Cynodontia. Many years ago Owen
recognised the remarkable mammalian characters in the Permian
and Triassic South Afvican reptiles, and though the Cynodonts
were so little known, he ventured to suggest that certain of the
Anomodonts were fairly closely allied, and perhaps ancestral, to
the Monotremes. Cope held much the same view. When the
very much more mammal-like Cynodonts were described by
Seeley, many recognised in this higher group the looked for Sauro-
Mammalia. Osborn has been the chief advocate of this opinion.
Seeley himself, though at, first inclining to it, afterwards came to
the conclusion that the Mammals were in no way nearly related
to the Cynodonts, but sprang from some unknown ancestor that
lived in Devonian or Silurian times.

If the Cynodonts are not nearly related to Mammals, the group
is still of great interest as showing a marvellous parallelism with
the Mammals in skull, teeth, girdles, limbs, and digits; but if,
as all recent work seems to indicate still more clearly, the mam-
malian ancestor was probably a Cynodont, the group becomes
vested with an interest altogether unique, and everything bearing
on it becomes worthy of the most careful study. I have fortu-
nately been able to examine every known skull, and in the present
paper I give the results of my researches. As the paper is
morphological rather than systematic, | propose to give a detailed
account of the skulls of only the principal Cynodont types, and
to consider more fully those points which seem to have a special
bearing on the question of mammalian descent.

Bauria.
(Pl. XLVI. figs. 6, 7, 8, and text-figs. 168, 169.)

Though Bauria cynops occurs in the same horizon as Cyno-
gnathus, it is the most primitive Cynodont at present known, and
may be regarded as the type of a distinct family which may be
called the Bauride.

As I have just recently, at considerable length, described the
only known skull of Bawria cynops, it will be unnecessary here
to do more than supplement that description in a few details
and to consider its relationships to the other known Cynodonts,
the Therocephalians, and the Mammals.

Further development and examination of the. skull has revealed
one or two points not previously noted. Under the nostril and
forming not only its floor but covering a considerable part of
the premaxillary is a large septomaxillary bone. The lachrymal
and prefrontal bones cannot in the specimen be clearly separated

896 DR. R. BROOM ON THE

from one another, but it is quite manifest that the lachrymal
is small and the prefrontal only moderate-sized. The nasal
extends well back and forms a broad suture with the frontal.
The frontals form the greater part of the interorbital region
and most of the supraorbital ridge. There are no postfrontals,
and the postorbitals are remarkable in forming only a postorbital
process and in not meeting the jugal to form a postorbital arch.
The jugal is slender and passes back nearly to the articular region.
The portion of the squamosal that supports the quadrate is well
developed, but the zygomatic portion is slender.

Text-fig. 168.*

Side view of the type and only known specimen of Bawria cynops. Since the speci-
men was first figured it has been considerably further developed at the British
Museum and by myself. The jaws are represented as closed. ‘The molar teeth
must meet one another as shown in the figure. When first described the teeth
were regarded as round, but further development shows that they are about
twice as broad in one diameter as in the other. Though the incisors are mostly
broken the impressions of the greater part of each is preserved, and the lower
must have met the upper as shown in the figure. The jugal arch is represented
in its central part only by the impression, but there is no doubt it must have
been practically as restored. It certainly did not meet the postorbital, which
is perfectly preserved on both sides. All the sutures shown in unbroken line
can be clearly made out.

The palate is as in typical Cynodonts, the secondary palate
being as well developed. The vomer, palatines, and pterygoid, so
far as can be seen, all are of the ordinary Cynodont type. The
lower part of the alisphenoid appears to be of the same type as in
higher Cynodonts, articulating with the basisphenoid and passing
out to the quadrate. It is just possible, however, that this
outward extension may be, as in the Therocephalians, entirely
formed by the pterygoid. The basisphenoid is unlike that of
either the Anomodonts, Therocephalians, or higher Cynodonts.
It is short, with a broad articulation for the basioccipital behind,
and a narrow articulation in front for the alisphenoid and
probably the vomer. From its under surface there passes down

* For explanation of the lettering in the: text-figures see p. 925.

SKULLS OF CYNODONT REPTILES. 897

a narrow deep median ridge, which is nearly as deep as the bone

is long. The basioccipital resembles considerably that of the

Anomodonts in having a pair of short postero-lateral processes
Text-fig. 169.

~Pmx.

757

Bi
ii

7,
ait,

FO.

Upper view of the skull of Bauria cynops.

which meet the basisphenoid. The condyle is unique (Pl. XLVI.
fig. 8). Itis asingle condyle, only partly divided into two by a
deep median groove. It is thus in type intermediate between the
condyle of the Therocephalian and that of the higher Cynodont.

898 DR. R. BROOM ON THE

The large foramen for nerves 1x., X., x1., and xi). lies by the side
of the basioccipital and in front of a bone which is probably part of
the opisthotic. Nerve xii. enters the foramen exactly as it does
in the higher Cynodonts. On the inside of the skull it has two
small distinct canals, which pass forwards and both unite with
the large foramen.

The bone which is supposed to be the stapes is shown in fig. 8
(Pl. XLVI). It is apparently a little displaced forwards.

The lower jaw has a fairly large surangular and angular, the
dentary being considerably in front of the articular region.

Taking all the characters into consideration, Bauria becomes
one of the most interesting intermediate types ever discovered.
Though an undoubted Cynodont, it retains many of the Thero-
cephalian characters. On the other hand, though on the whole
it is less mammal-like than the higher Cynodonts, it has some
mammalian characters which the others have lost.

The following are Therocephalian characters usually lost in
Cynodonts but retained in Lawria :—-

1. Large septomaxillaries forming part of the facial surface.

2. Moderate prefrontals.

3. Large frontals forming most of the interorbital region.

4. Feeble zygomatic arch.

5. The two occipital condyles so imperfectly separated as to
represent practically a single condyle.

6. Large size of angular and surangular.

7. Shape of the articular.

8. Simple condition of the molar teeth.

In the following characters Bawria is nearer to the mammalian
ancestor than are the higher Cyiodonts :—

1. Large size of septomaxillaries and development on face.
A somewhat similar condition is found in primitive -
Multituberculata (e. g. Tritylodon), also in Monotremata,
as shown by Gaupp in Lchidna embryo.

. Large frontals.

Complete loss of parietal foramen,

Absence of postorbital arch.

5. Simple condition of molar teeth.

bm Gols

Nythosaurus,
(Text=fig. 170.)

The type of Galesaurus planiceps is a somewhat céushed skull
with the bones in an unsatisfactory condition for showing sutures.
No second specimen of Galesaurus has ever been discovered.
In 1876 Owen des¢ribed an imperfect skull as Vythosaurus lar-
vatus. In 1887 he described another skull in fairly good preserva-
tion which he believed to be another specimen of Galesaurus.
Seeley in 1894 showed that this supposed second specimen of

SKULLS OF GYNODONT REPTILES. 899

Galesaurus differed greatly from the type, and gave it the name of
Thrinaxodon liorhinus.

There is, in my opinion, not the least doubt that Seeley was
right in regarding the 1887 skull as belonging to a very different,
animal from the 1859 one. In fact it seems strange that any one
should ever have thought them the same. Galesaurus has a

dental formula apparently of 1. 2 c. a m. ae the 1887 specimen
G. , m. q Tn the 1859 specimen 10 molars
occupy 20 mm.; in the 1887 specimen 7 molars occupy 20 mm.
But while the two supposed Galesaurus specimens represent
different genera, two other imperfect specimens in the British
Museum show that the 1887 specimen is the same animal as was
described in 1876 as Nythosauwrus larvatus. Hence the well-
known skull which is figured in various text-books as Galesaurus
must in future be called Vythosaurus.

Nythosaurus is a much higher type than Bawria: but though
it comes fairly close to the higher Cynodonts such as Cynognathus,
it should, I think, be taken as the representative of a distinct
family, the Galesauride. From the various specimens in the
British Museum it is possible to make an almost complete restora-
tion of the skull.

has a formula of 1.

Text-fig. 170.

Side view of the skull of Nythosawrus larvatus. The drawing is mainly that of the
best preserved specimen in the British Museum, compared with the other
specimens and slightly restored from them. ‘The teeth are represented in the
mature condition.

The septomaxillary though smaller than in Bauria still appears
on the face. The nasal is large and very broad at its upper end.
The lachrymal is large, and though the prefrontal is only of
moderate size, it joins with the postorbital and completely shuts
out the frontal from the orbit. The postorbital forms with the

900° DR. R. BROOM ON THE

jugal a rather feeble postorbital arch. The zygomatic arch is
formed by the jugal and the squamosal. ‘The jugal extends
nearly back to the articular region, and the squamosal nearly
forward to the base of the postorbital arch. The squamosal is
not unlike that of Bauwria, but the zygomatic portion is much
better developed ; so that as regards the squamosal Vythosaurus
is intermediate between Bauria and Cynognathus. The quadrate
is of the same type as in the better known Cynognathus.

The palate, so far as known, agrees fairly well with the Cyno-
gnathus type, and the occipital condyie is double.

The lower jaw has a large dentary, but there is no trace even
of a condylar process. The angular and surangular are fairly
large and still resemble considerably the Therocepbalian type.
The articular also resembles that of the earlier rather than that
of the later types.

Nythosaurus is perhaps the most mammal-like of the known
Cynodonts. The zygomatic arch is exceedingly like that of most
primitive mammals, and if the prefrontal and postorbital bones
were lost and the internasal process of the premaxilla aborted
there would be nothing left in the side view of the skull to
distinguish it from that of a mammal. The lower jaw with its
fairly large angular and surangular is still much less like the
mammalian condition than what we see im the higher Cynodonts,
and the articular is of the same primitive type seen in Lauria.

The dentition though very primitive is considerably more
1 7
3) D mM.

comes very near to that of the typical mammal, and that of

highly evolved than in Baeria. The TOLEDO, ty Sy Ce

7?

o 4) 1 i2
Galesawrus, i. 5, C- 72 M. 7, 1S near that of the ancestral mammal.

The difference in the teeth in some of the specimens of Vytho-
saurus is apparently due to the fact that in some the teeth belong
to the first set and in ethers to the second.

Cynognathus,
(Pl. XLVI. figs. 1 & 2, and text-figs. 171, 172.)

The genus Cynognathus is known by the very fine skull of
(. crateronotus in the British Museum, the type skull of C. platy-
ceps in the Albany Museum, a fairly good skull of C. berryiin the
S. African Museum, and three or four less satisfactory specimens.

Seeley has given a fairly full account of the skull of Cyno-
gnathus crateronotus, but unfortunately a number of his figures are |
so indifferently reproduced that they convey no more to the
student than does the plaster cast. And further, while most of
his determinations are correct, he unfortunately suggests so many
alternative possibilities that the morphologist is left compara-
tively helpless.

The figure given by Seeley of the side view of the skull of
C. crateronotus gives an excellent idea of the general form of the
skull and of the structure of the temporal region, except that the

SKULLS OF CYNODONT REPTILES. 901

supposed perforation in the zygomatic arch is, in my opinion, not
a natural feature and has been produced post mortem. The side
view which I give of the skull of C. platyceps is fairly similar,
except that the skull is here much broader and flatter, and in
this species, at least, there is no trace of an opening in the
zygoma.

The snout of Cynognathus differs from that of Sauria and
Nythosaurus chiefly in the fact that the septomaxillary does not
appear on the face, though it can be readily seen within the
nostril.

Text-fig. 171.

hip,
Vi Less
MI

Side view of the skull of Cynognathus platyceps. With the exception of the
tront of the snout the drawing is made trom the very fine type skull in the
Albany Museum. The front of the snout is from the specimen described by
Seeley as ? Cynognathus leptorhinus. As I showed some years ago, this is
unquestionably the snout of a nearly full-grown specimen of Cynognathus
platyceps.

The premaxilla is relatively rather larger than in Vythosaurus,
while the maxilla is about equally well developed in the two
genera. The canine is, however, much further forward in Cyno-
gnathus than in the smaller genus.

The nasal bone is fairly similar in the two genera, being broad
both in front and behind and narrow in the middle.

The lachrymal extends further forward than in WVythosaurus,
the portion showing on the face being nearly as large as the
orbit.

The prefrontal is a long narrow bone which forms the greater
part of the upper margin of the orbit, and by meeting with the
postorbital completely shuts out the frontal from the orbital
margin.

The frontals are each about four times as long as_ broad.
In front they meet the nasals. Laterally they are in contact
with the prefrontals and postorbitals. Posteriorly they taper
away to narrow points, which mect the anterior ends of the
parietals.

The postorbital is a large triangular bone. In front it meets
Proc. Zoou. Soc.—1911, No. LXII. 62

902 DR. R. BROOM ON THE

the prefr ontal and forms part of the orbital margin. Externally
it gives a large articulation to the jugal and a small articulation
to the squamosal, Internally it overlaps the parietal for only a
very short distance.

The jugal is relatively considerably larger than in Vythosaurus.
Anteriorly it meets the maxilla and jachrymal, but extends further
forward thanineither Vythosaurus or Bauria. Immediately below
the orbit there is a small but very distinct tubercle. The
ascending process of the jugal is unusually broad and forms a
large articulation with the postorbital, the two together forming
a very strong postorbital arch. The posterior portion of the jugal
extends to the articular region.

The parietal is a narrow bone which forms a low median crest.
There is a small pineal foramen.

The squamosal is the largest bone in the skull, with the excep-
tion of the dentary. The inner and posterior portion forms
nearly the whole of the back wall of the temporal fossa, and has
a large articulation with the parietal. This back portion of the
squamosal is very thin and is closely united with the flat upper
expansion of the opisthotic. The squamosal forms the outer wall of
the lateral occipital foramen. Inferiorly it meets the exoccipital
and on passing outwards supports the small quadrate. The
zygomatic portion is much larger than in Vythosaurus, articulating
with the whole of the upper side of the posterior limb of the
jugal and meeting the postorbital. There is a deep groove along
the middle of the back part of the zygomatic portion, whieh
curves downwards and inwards, and, most probably, the pos-
terior part was for the support of the external auditory canal.

The occiput is best known from the specimen of Cynognathas
berryt in the 8. African Museum, which though imperfect shows
the sutures very distinctly. The lar “ge occiput figured by Seeley*
(p. 130) and doubtfully referred by him to @. hoe yi, 1S, IN my
opinion, considerably too large. It also differs from the known
occiput of C. berryi in the ‘shape of the foramen magnum, the
slope of the exoccipitals, and the moulding of the interparietal
region, Itis pretty clearly not the occiput of C. crateronotus,
while it is much too lar ge to be that of C. platyceps. As it thus
seems to belong toa new species, it may appropriately be named
Cynognathus seeler Ye

The occiput proper (Pl. XLVI. fig. 2) is made up of seven
bones, or, if we include the squamosals, nine. About a quarter
of the occiput proper is formed by a median bone, which is
apparently the interparietal. I have not seen any specimen which
enables me clearly to differentiate this bone from the parietal in
front, but, judging by analogy and by the direction of the fibres
of the bone, the probability seems much in favour of its being a
distinct interparietal.

On either side of the interparietal is articulated a large bone,
which is apparently the opisthotic. Its occipital por tion is com-
paratively thin and to a considerable extent covered in front by

* Phil. Trans. Vol. elxxxvi. B. (1895).

SKULLS OF CYNODONT REPTILES. 903

the squamosal and parietal. Inferiorly it articulates with the
exoccipital and to a large extent with the supraoccipital, It

Text-fig. 172.

Upper view of the skull of Cynognathus platyceps-
From the same specimen as text-fig. 171.

forms the upper and inner wall of the lateral occipital foramen,
and passes well forward below the edge of the parietal and appears
to articulate with the large prootic.

62*

=!

904 DR. R. BROOM ON THE

The supraoccipital is completely anchylosed to the exoccipitals,
but its probable limits are indicated in the figure given, [mme-
diately above the foramen magnum is a small ‘rounded bony knob.

‘The exoccipitals form the greater part of the oceipital condyles,
whieh are remarkably mammal-like. Above each condyle is a
narrow fissure or groove leading into the foramen magnum, but
it seems too small to have been for the transmission of either an
artery or a vein, and most probably it was not for a nerve.
Possibly ib is merely owing to the elevation of a portion of the
bone for the attachment of a ligament. The exoccipital passes out
aconsiderable distance under the lateral foramen and articulates
with the squamosal.

The quadrate is well shown in Seeley’s figure 8. It is a
relatively small bone, which is tirmly fixed on to the squamosal by
the main part being in front and two long processes being behind
its lower projection. Referring to the posterior delicate processes
Seeley says: “ I am unable to attirm that they represent auditory
ossicles.” They are unquestionably parts of the quadrate and
have nothing to do with the auditory function. The quadrate
forms the greater part of the articular surface for the lower jaw,
but not the whole of it, part of the squamosal also forming
a portion of the articulation. This is particularly interesting in
view of the fact that in the Monotremes the lower jaw hinges
directly on the squamosal bone.

The palate of Cynognathus is nearly wholly known, the only
points concerning which we are still ignorant being the relations of

the bones round the anterior palatine foramina, the nature of the
middle part of the basicranium, and the relations of the palatine
and pter ygoid to the jugal.

‘There is a large secondary palate formed by the maxille and
palatines exactly as in Mammals. In a recent paper Seeley
describes what he believes to be teeth on the palate of
Cynognathus. I have, however, had an opportunity of examining
Seeley’s specimen and believe the supposed teeth to be merely
irregularities of the bony surface, possibly pathological. Certainly
in the other specimens I have examined there is no trace of
anything like teeth. The hard palate ends in the middle line
opposite the front of the third last molar.

The pterygoids are large and have well- -developed pterygoid
processes, which lie close along the inside of the jaws as in reptiles
generally. There is no transpalatine or ectopterygoid bone.
Instead of, as in most reptiles, the pterygoid having a posterior
process which extends to the quadrate, it here ends near the
middle of the inner wall of the temporal fossa, the posterior
continuation which looks like pterygoid being really part of the
alisphenoid bone.

The vomer is a large median bone which posteriorly les between
the two pterygoids. In front it forms a vertical plate which
supports the secondary palate exactly as in Mammals and extends
to near the front of the snout.

SKULLS OF CYNODONT REPTILES. ~ 905

The alisphenoid bone is one of the most interesting bones in
the skull. It isa large flat bone which extends from the pterygoid
below to the parietal above. It is well seen in the type skull of
Cynognathus crateronotus, but even better in the Capetown
specimen of C. berryi (P]. XLVI. fig. 1). The upper portion of
the bone is irregularly quadrilateral. The upper side articulates
with the parietal, and the posterior with what I believe to be the
prootic. Between the alisphenoid and the prootic are two large
oval foramina. At the posterior and lower corner the alisphenoid
is continued as a slender bone to the quadrate. At its anterior
and lower corner it meets the pterygoids and clasps the basi-
sphenoid. There appears to be an opening into the brain-cavity
between the base of the alisphenoid and the basisphenoid.

The basisphenoid is clasped by the alisphenoids and meets the
basioccipital posteriorly.

The basioccipital is a small bone lying behind the basisphenoid.
It forms the middle part of the occipital condylar region. On
each side’ there is a large round foramen which is pretty certainly
the foramen for the exit of nerves ix., x., xi., and xii.

Between the outer part of the basioccipital and the quadrate
there stretches a rounded pillow-like bone concerning which there
may be some difference of opinion. I believe it to be the
stapes, for reasons which will be stated later.

The lower jaw is remarkable for the great size of the dentary,
which posteriorly nearly reaches the articulation. Elsewhere I
have dealt at some length with the structure of the jaw. The
splenial is long and slender. The surangular and angular are also
feeble splint-like bones. The articular is fairly well developed but
short. I cannot satisfy myself that there is a distinct coronoid
bone as is stated by Seeley.

The only points in which the Cynognathus skull is nearer to
the mammal than that of Bawria and Nythosaurus are: (a) the
closer approach of the jugal to the articulation, (6) the greater
development of the dentary, (c) the greater reduction of the
angular and surangular, and (d) the more mammal-like occipital
condyle. On the whole it is not so near the mammalian ancestor
as either Bauria ov Nythosaurus.

The dental formula appears to be i. 4, ¢. * 3

7 Col ay) Miler ge

Trirachodon.
(Text-figs. 173 & 174.)

Trirachodon is best known by the type skull which is in the
Albany Museum. Though the skull is immature and much
crushed it is practically perfect. Two or three other known skulls
though imperfect show the uncrushed condition of the greater part
of the adult skull.

The premaxilla is smaller than in Cynognathus, not meeting the
nasal behind the nostril, at least not on the face.

906 : DR. R. BROOM ON THE

The septomaxillary is entirely in the nostril, forming no part of
the face.

The maxilla is long and extends far back below the orbit.
The snout is fairly broad at the root of the canine and along the
upper part of the maxilla, but is much narrowed in the molar
region. There are two foramina for the maxillary branch of
nerve Y.

The nasal is moderately broad in front, narrow in the middle,
and very broad behind.

The lachrymal is small, but forms most of the anterior wall of
the orbit.

Text-fig. 173.

Ju-

\

Zz_
gg

ts
/

/
Y

Side view of the skull of Trirachodon kannemeyeri. The drawing is chiefly founded
on the crushed and immature skull which forms the type. Two mature
and uncrushed but imperfect skulls in the Albany Museum and a good snout
in my own collection have made it possible to correct the crushing of the type
and completely restore the skull in the adult cqndition.

The prefrontal is about twice as Jong as broad, and forms most
of the upper margin of the orbit; it unites, as in Cynognathus,
with the postorbital, completely shutting out the frontal from the
orbital margin.

The frontal is fairly like that of Cynognathus, but it does not
extend so far forward. Posteriorly, as in the former genus, it
tapers away between the postorbitals.

The postorbital is more like that of Vythosaurus. It forms the
upper third of the postorbital arch, uniting with the jugal. It
extends backwards on the side of the parietal a little beyond the
pineal foramen.

The parietal is like that of Cynognathus, but the pineal foramen
is much smaller.

The jugal is, on the whole, like that of Cynognathus. It has,
however, the inferior process much better developed. It forms
the lower and posterior half of the orbital margin.

The squamosal differs from that of Cynognathus in not meeting
the postorbital, but ending in front above the jugal, very much as

SKULLS OF CYNODONT REPTILES. 907

in Vythosawrus. As the back of the skull is relatively narrower
than in Cynognathus, the posterior part of the squamosal differs
considerably in contour.

Text-fig. 174.

HH
) <a Th
‘ BRASS Nes
ANS RSSET
ASS
BSR TA

EA

THT
Lsbeeas
as:

ef
Lif

7
2.

none,

ere,

NS a we

Upper view of the skull of Tvirachodon kanneneyert.
From the same specimens as fig. 173.

The quadrate, so far as can be seen, is small but not unlike the

better known types.
The stapes is a very slender straight bone about the thickness

908 DR. R. BROOM ON THE

of a pin, and having similar relations to the stapes in Cyno-
gnathus.

The occiput is not well known, but is apparently very similar
to that of Cynognathus.

The lower jaw has a very large coronoid process and a condylar
process which nearly reaches the articulation. The articular,
angular, and surangular are on the whole very similar to those of
Cynognathus.

One of the most striking points of difference from Vythosaurus
and Cynognathus is in the structure of the molar teeth, which
have flattened tops, and the lower molars instead of passing inside
of them, as in these other genera, meet them much in the same
way as do the molars in Mammals.

The dental formula is the same as in Oynognathus, viz.:

» 4 1 9
Tas) © ap 0S 50

Diademodon and Gomphognathus.
(Pl. XLVI. fig. 9, & text-figs. 175-178.)

Diademodon resembles Gomphognathus so closely, differing only
in size and in the number of molars, that there is some reason for
suspecting that Diademodon may be animmature Gomphognathus.
If this turns out to be the case, the genus must take the earlier
name Diademodon. The following description of a skull is based
on a beautifully preserved skull in the British Museum, which
may be called Gomphognathus minor, but which unfortunately has
lost the snout, on a fairly good skull of D. mastacus in the South
African Museum, on two other good skulls of Gomphognathus in
the British Museum, and on the type skull of Gomphognathus
kannemeyert in the Albany Museum. As the result of the
examination of this very fine material, the Gomphognathus skull
is better known even than that of Cynognathus, and almost as
well as that of the living Ornithorhynchus.

The premaxilla is fairly large, but, as in all other Cynodonts,
considerably overlapped by the front of the maxilla, It has a
strong internasal process. and forms the anterior and most of the
lower border of the nostril. It has a very considerable palatal
development, the two bones meeting in the middle line behind the
anterior palatine foramen.

The septomaxillary is well developed, lying along the greater
part of the outer wall of the nostril. Most of it is within the
nostril, but a small part of the upper end appears on the face.

The maxilla is not unlike that of Zrirachodon. The maxillary
branch of nerve v. has two foramina. On the upper part of the
maxilla close to the nasal is a small oval depression, presumably
for the lodgment of a gland.

The nasal is narrow in front but broad behind. The nostrils
look more upwards than in any of the previously described
Cynodonts, and the nasal passes forward between them to a
narrow process.

SKULLS OF CYNODONT REPTILES. 909

The lachrymal is of fair size. In Gomphognathus minor it
forms the front of the orbit and much of the inner wall. In
Gomphognathus polyphagus ib is considerably larger, extending to
part of the upper margin.

Text-fig. 175.

Tiki

I

Side view of the skull of Gomphognathus minor. With the exception of the front
half of the snout and the lower jaw, the drawing is from the beautifully preserved
type in the British Museum. The front of the snout is restored from a specimen
ot Gomphognathus kannemeyeri 1m the British Museum, and the lower jaw 1s
from the lower jaw of the type of G. kannemeyeri in the Albany Museum.
Both these latter are slightly modified to fit the skull of Gomphognathus
minor.

The prefrontal forms most of the upper margin of the orbit.
In Gomphognathus polyphagus it is considerably smaller than in
G. minor, owing to its being encroached on by the larger nasal and
lachrymal. By uniting with the postorbital it completely shuts
out the frontal from the orbital margin.

The frontal is relatively small, the two together forming
only about one-third of the interorbital space. As in all the
other Cynodonts except Bauwria and Sesamodon, the frontals
posteriorly taper away to a point between the postorbitals.

The postorbital is a fairly large bone with an external limb
which meets the jugal, forming the postorbital arch, and a posterior
process which les along the parietal. In G. polyphagus the
postorbital bar is relatively slenderer than in G. minor.

The jugal is very large. It is essentially similar to that of
Trirachodon. The inferior process is much larger and the
posterior extension much deeper. It passes backwards some
distance behind the plane of the quadrate.

The parietal is small and there is a small pineal foramen.

The squamosal is extremely large. The inner portion
articulates with the parietal. The outer and anterior portion
lies above the jugal, forming with it a powerful zygomatic arch.
Inferiorly the squamosal supports the small quadrate. The
peculiar shape of the bone can best be understood from the figures.

910 DR. R. BROOM ON THE

The occiput is very similar to that of Cynognathus, but the
limits of the various elements are less satisfactorily known. The
condyle is double, but is relatively smaller and less mammal-like
than in Cynognathus.

The palate is beautifully shown in three of the British Museum
specimens,

Text-fig. 176.

Upper view of skull of Gomphognathus minor.
The snout restored from Gomphognathus kannemeyeri.

In the anterior palatine opening there is a pair of narrow bones
showing what I suggested a good many years ago were probably
prevomers. Only one specimen is known in which they are shown.
They are evidently not parts of the premaxille, and as they are in
position exactly corresponding to the prevomers of Ornitho-
rhynchus, | am still of opinion that they correspond to the paired
vomers of most reptiles and the prevomers of Platypus and of
Miniopterus.

SKULLS OF CYNODONYT REPTILES. 911

The secondary palate is almost typically mammalian, the maxillee
and the palatines having the same relations as in the mammal,
The palatine besides forming part of the secondary palate curves
round inside of the maxilla, forming the outer wall and part of
the roof of the posterior nares. There is a posterior palatine
foramen situated exactly as in Mammals.

Text-fig. 177.

IMG XXI, XI.

Under view of skull of Gomphognathus minor.
The snout restored from Gomphognathus kannemeyeri.

The vomer is large and, as in typical Mammals, it forms the
median support of the basicranial axis from the sphenoidal region
to near the front of thenose. It forms about one-third of the roof
of the pharynx, and two small tubercles on its posterior part are
probably for the attachment of pharyngeal muscles. In the
region of the secondary palate the vomer forms for a considerable
distance a median support.

The pterygoid is of moderate size, but much smaller than in any

912 P DR. R. BROOM ON THE

other reptilian group. In front it meets the jngal and the
palatine and lies along the outer side of the vomer. It forms a
large descending pterygoid process. Posteriorly it lies against the
vomer and terminates by meeting the alisphenoid as in
Cynognathus.

Text-fig. 178.

=
SS

Posterior view of skull of Gomphognathus minor.

The basioccipital, basisphenoid, and exoccipital are all
apparently very similar to those bones in Cynognathus. There
is a large foramen for the exit of nerves ix., x., xi., and xi.
Outside of this foramen, but further from the base, is another
foramen which I believe to be the fenestra ovalis, but this 1s less
certain than the nature of the larger foramen, about which there
is no doubt.

The alisphenoid in all its relations is very similar to that in
Cynognathus. At its lower end near where it meets the pterygoid
is an opening which leads into the brain-cavity, and most probably
it was by this opening that the internal carotid artery entered

‘the cranium.

The quadrate is relatively rather smaller than in Cynognathus,
but is fixed into the squamosal bone in a very similar fashion.
Tt clasps the lower margin of the bone, and posteriorly it has two
processes which fit into grooves. In the type specimen of
Gomphognathus kannemeyert, where the articulars do not fit on to
the quadrates, I thought the quadrates had been displaced, but I
am rather inclined to think it is the articulars, as the London
specimens show that the quadrate is not likely to be readily
disarticulated.

In the median section of the skull (Pl. XLVI. fig. 9) a number
of most interesting features are revealed. In the posterior
cranial region there are seen the foramina for the exit of a
number of the cranial nerves. Close to the occipital condyle
are two small foramina for nerve xii. These after passing a
short distance through the bone open into the large foramen
lacerum posticum. This large foramen is also situated well back
and doubtless transmitted also nerves ix., x., and xi. In front

SKULLS OF CYNODONT REPTILES. 913

of this foramen jugulare is the prootic bone, which appears to
have two foramina in it. The posterior I believe to be for
nerve vill., aud the anterior for vii. Between the prootic and
the large alisphenoid is seen the opening for probably both the
2nd and 3rd branches of nerve v. _ Immediately below this
foramen, the prootic sends a sharp bony process upwards, inwards,
and forwards. Probably it lay on the inner side of the Gasserian
ganglion.

The basisphenoid is a large bone, along the front of which lies
the back part of the vomer. There is no orbito-sphenoid and no
presphenoid bone.

The vomer passes from the basisphenoid to about the middle of
the hard palate. Along its dorsal surface ran the cartilaginous
cranial axis, against which the grooved upper surface of the vomer
fits. At the front of the vomer the median cartilage is ossified,
and the bone is apparently the homologue of the mammalian
mesethmoid. In the figure given of the median section the
prevomer (P.vo.) and the septomaxillary (Sma.) are largely
hypothetical.

Sesamodon and Alelinodon.

(Pl. XLVI. figs. 3, 4, & 5, and text-figs. 179, 180.)

These two allied genera, which are both unfortunately very
imperfectly known, stand at present by themselves some distance
apart from the other Cynodonts. Each genus is known only by
a single specimen, which in the case of Scsaimoden brownt 18 only
fair, and in the case of Melinodon simus ver ypoor. Still the interest
attaching to the specimens is so very great that it is necessary to
figure them as fully as possible.

Though the only known skull of Sesamodon browni is very badly
weathered and considerably crushed, it is fortunately possible to
restore the external appearance with much certainty. In fact the
only points in the external anatomy that remain in doubt are the
articular region, the middle of the occiput, and parts of the jugal,
frontal, parietal, and squamosal bones.

The premaxilla is not very satisfactorily preserved, but presents
no unusual features.

The septomaxillary is apparently fairly similar to that of
Nythosaurus, appearing on the face to a considerable extent.

The maxilla is large and resembles to some extent that of
Trirachodon, while in other respects it is nearer to Vythosaurus.
The canine is relatively smaller than in either Zrirachodon or
Nythosaurus, but is situated, as in Vythosaurus, far back from
the front of the bone. The molars, with the exception of the Ist
which is small, form a uniform series, and the maxillary bones are
much approximated in the molar region, as in Zrirachodon.

The nasal resembles on the whole that of Bauria. The nostril
is directed mainly forward and the nasal to some extent over-
hangs it. The bone is moderately broad throughout its whole

914 DR. R. BROOM ON THE

length, but is chiefly remarkable in being narrower behind than
in front. In all other Cynodonts except Lauria the nasal is, as in
Marsupials, much broader posteriorly.

Text-fig. 179.

Side view of skull of Sesamodon browni. Somewhat restored from the only known
specimen, which forms the type. Ihe whole of the preorbital portion of the
skull except the front of the premaxilla is preserved in the specimen, though the
bone in parts is weathered off, leaving only the impression. Though the teeth
are imperfect, remains or impressions of all are present, so that the full dentition

_can be restored with much certainty, the only doubt being the exact length of

“the incisors and canines. The orbit and the temporal fossa are satisfactorily
preserved, and the squamosal is fairly well preserved on the right side of the
skull. The lower border of the jugal is unknown. Both mandibles are in
position but much weathered. The horizontal ramus ts fully known, but much
of the ascending ramus is lost. As, however, the top of the coronoid process 1s
preserved in position, the greater part of the dentary can be restored with
certainty. The condylar process is badly preserved. A considerable part of
what is believed to be the angular and prokably part of the articular are
preserved. As the position of the glenoid cavity is known, the general shape of
the back of the jaw can be restored with some probability.

The lachrymal is small and completely separated from the nasal
by the prefrontal. In this, Sesamodon again agrees with Lauria
and differs from all other known Cynodonts.

The prefrontal is much larger than the lachrymal and meets the
frontal, nasal, maxilla, and lachrymal as in auria.

The frontal is almost completely lost from the specimen, but
just sufficient of the impression of the bone is left to show that it
reached the orbit as in Bauria. In this also, Sesamodon differs
from almost all other known Cynodonts.

The postorbital is fairly large but slender. It forms with a
small part of the jugal a complete postorbital arch. It only
extends backwards on the parietal a very short distance.

The parietal is completely lost except just sufficient to show
the width of the bone.

The jugal is not well preserved except in the upper part. . It
forms the whole of the lower orbital margin and part of the

SKULLS OF CYNODONT REPTILES, 915

postorbital arch. In the drawing given the lower margin is
entirely hypothetical. The posterior or zygomatic portion of the
bone is very short.

Text-fig. 180.

Upper view of skull of Sesamodon browmi, restored from the type. From the specimen
the whole of the frontals, parietals, and most of the occiput have weathered away,
so that these parts are unknown. "As, however, the prefrontals, postorbitals, and
the margins of the orbits and temporal fosse are preserved, the only points
that are left in doubt are the position of the fronto-parietal suture, whether
there is a parietal foramen, and the nature of the condyles.

The squamosal is sufficiently well preserved to show its main
features, and it is seen to be unlike that of any other Cynodont.
As in Bauria it is much smaller than in the higher types. It
articulates, as in other forms, with the parietal, but on passing
outwards it has not, asin Bawre ‘1a, a poster ior ridge. The zygomatic
portion is much Shomhen than in Bawria or any other known type,
but it is relatively fairly deep.

What appears to be the quadrate is a small flattened bone which
I have shown in the figure, but the parts are crushed and some-
what displaced, and it is impossible to speak with certainty of the
condition.

916 DR. R. BROOM ON THE

The occiput is badly preserved, but is remarkable for the great
lateral extension of the exoccipital, which passes out behind and
below the squamosal. Owing to some degree of crushing it is
difficult to be quite sure of the relations of the external end of
the exoccipital, but it certainly extends much further out than in
any other known Cynodont.

The palate, so far as preserved, differs considerably from that of
the typical Cynodonts. There is asecondary palate of the ordinary
type, but the pterygoid appears to have a larger pterygoid process
than usual. Behind the pterygoid process there isa fan-like bony
expansion which passes backwards and outwards towards the
articular region. It looks as if it might be all pterygoid, but
owing to the crushed and weathered condition of the specimen it
is impossible to be sure. Possibly, as in the typical Cynodonts, it
is part of the alisphenoid.

The structure of the lower jaw cannot be satisfactorily made
out. The dentary has a very large coronoid process ; in fact the
coronoid process is as large as the horizontal ramus. There also
is some evidence of a condylar process. The articulation is
apparently, mainly at least, formed bya rounded articular supported
by possibly an angular and surangular. There is a well-developed
splenial bone.

m

Pages A:
The dental formula is 1. 3, ¢. 7, m. ;

Melinodon is closely allied to Sesamodon and pretty certainly
belongs to the same family. The teeth are of the same type, but
relatively much smaller. The specimen is so imperfect that it is
impossible to make much of the skull. I have figured it as
preserved (Pl. XLVI. fig. 3).

Sesamodon resembles Bauria and differs from the other
Cynodonts in the following characters :—

Uae

1. The nostril is directed more forwards than outwards.

2. The nasal is not widened posteriorly.

3. The prefrontal is larger than the lachrymal and prevents
the lachrymal from meeting the nasal.

4. The frontal forms part of the orbital margin.

5. The postorbital arch is feeble: incomplete in Bauria.

6. The molars show no sign of cusps.

In the following characters Sesamodon comes nearer the
Mammals than any of the other known Cynodonts :—

1. An articulation for the lower jaw which permits of some
degree of antero-posterior movement.

2. The lower canine lies outside the edge of the maxilla when
the jaw is closed.

In addition to the mammalian characteristics peculiar to
Sesamodon, it combines most of those mammalian characters
seen in Bauria with most of those found in the other higher
Cynodonts.

SKULLS OF CYNODONT REPTILES. 917

Peculiarities of the Mammalian Skull, apparently derived
from a Cynodont Ancestor.

Most of the bones of the mammalian skull have their homo-
logues in the Labyrinthodont skull, but they are also to be found in
the skulls of most reptiles. There is, however, no close resemblance
between the mammalian and the batrachian bones, and in many
cases the differences in cranial structure are so great that the
gap between the mammal and any known batrachian must be
enormous. When we examine the Cotylosaurian skull, we find that
the resemblance to that of the mammal is still remote, but any
little resemblance there was in the Labyrinthodoent is here
increased, while there are many mammal-like characters not seen
in the lower type.

The Pelycosaurians of the Lower Permian are so much more
mammai-like than any of the lower forms that, notwithstanding
their remarkable specialisations, one cannot help feeling, as Cope
felt, that here were forms fairly near to the remote mammalian
ancestor.

The Therocephalians and Anomodonts of the Middle Permian
times are in essentials still more mammal-like. For the first time
we get a dentition clearly divided into incisors, canines, and
molars ; for the first time we get a lower Jaw witha dentary which
has a large coronoid process. We get a zygomatic arch formed on
the mammalian type, and we lose for the first time the quadrato-
jugal. We also get most marked mammalian characters in the
posteranial skeleton.

The Therocephalians survived into Upper Permian times, but
hitherto they have not been found in Triassic beds. In Upper
Triassic times their place was taken by the Cynodonts. Though
the gap between the Therocephalians at present known and the
Cynodonts is very considerable, the primitive Cynodont Bawria is
to some extent a connecting link.

Almost all the characters in which the Cynodont skull differs
from the Therocephalian are characters which are met with in
Mammals. Of these the most noteworthy are :—

1. Formation of a secondary palate.

2. Vomer very large, extending forward as a support to the
secondary palate.

Great reduction or complete loss of prevomers (Lauria).

Loss of the postfrontal bone.

Great reduction or loss of the pineal foramen.

Two occipital condyles.

Reduction of the quadrate.

A large alisphenoid bone instead of the homologous rod-
like ‘‘ epipterygoid” or columella cranii of the Thero-
cephalians and Anomodonts.

9. Pterygoids not extending back to the quadrates, the
posterior extension being replaced by the alisphenoids.

10. Reduction of the angular and surangular, and greater

development of the dentary.

Proc. Zoou. Soc.—1911, No. LXITI. 63

CoN See Se

918 DR. R. BROOM ON THE

The presence of this large number of mammalian characters in
the Cynodont skull, and the absence of any in either skull or
skeleton that might not have been expected in the mammalian
ancestor, make the case very strong for the mammalian ancestor
having been a C ynodont. But the evidence becomes even stronger
when we find that most of the peculiarities, even minor
peculiarities, of the mammalian skull have light thrown on them
by the condition of affairs in the Cynodont skull.

Let us consider some of the more remarkable characters of the
mammalian skull in the light of our knowledge of the Cynodont.

Premaaillary.

One of the most striking peculiarities of the mammalian skull
is that the nostrils are separated only by cartilage, so that if the
cartilage be removed the nostrils are united. In most reptiles, in
birds and amphibians the nostrils are divided by an upward and
backward process of the premaxilla, the internasal process. As it
is present in Cotylosaurs, Dromasaurs, Pelycosaurs, Therocepha-
lians, Anomodonts, and even Cynodonts, one might fancy that
here was evidence against the Cynodont ancestry. But there is
good reason to believe that the early Mammals retained the
internasal process and that it was only lost after the Mammals
were well established.

In both Ornithorhynchus and Hehidna the young animal has an
internasal process developed on the premaxilla almost exactly as
in reptiles. The fact that it is retained as a support to the
caruncle or egg-tooth in no way invalidates the conclusion that it
is the reptilian internasal process that has been retained. For
there cannot have been a time when there was acaruncle without
a support, and thus the internasal process must be as old as the
caruncle. As we may be pretty certain that the mammalian
ancestor was oviparous, we may safely conclude that the internasal
process is not a neomorph, but the reptilian structure handed on,

In Zritylodon there is an imperfect but distinct little internasal
process. The only known specimen is too imperfect to enable me
to say whether it formed a complete though slender process which
joined with the nasals. Even if it did not in the adult, it is
vather probable that it did in the very young animal, since
Tritylodon is so much more primitive than the marsupial that not
improbably it was oviparous.

Tn the skulls of young Diprotodonts (e. g. J/acropus) a rudiment
of the internasal process is usually present. And in the young
Trichosurus at birth the internasal process, as I recently pomted
out, can be traced right round in front of the nose. In the very
young marsupial, the nostrils are entirely lateral and wide apart,
and the nasal cartilages pass round in front of each, leaving a
sulcus in the middle line between tke two. The premaxillaries
send up short processes along the sulcus, but from the ends of the
processes two strands of condensed but unossified cells can be easily

SKULLS OF CYNODONT REPTILES. 919

traced round to the top of the snout. Were these tracts ossified
we would have a condition practically similar to that of the
Cynodont.

Septomaxillary.

Kitchen Parker many years ago recognised this as a distinct
membrane-bone in the Lizards and Snakes, but it is only recently
that much attention has been paid to it, chiefly as the result of the
work of one or two paleontologists in Europe and America. The
bone is not known in Labyrinthodonts and probably does not
occur in the Amphibia (the supposed septomaxillary of Xenopus
being probably not homologous). We find it, however, in the very
earliest true reptiles, and we can trace it on the one hand through
numerous members of the Diapsida, and on the other through
most of the mammal-like Reptiles on to Mammals.

A septomaxillary has been found in Pareiasaurus, Pariotichus,
and Procolophon. In these primitive genera it is mainly within
the nostrils, and probably fulfils its main function as a roof to
Jacobson’s organ.

When we come to the mammal-like Reptiles, we find it in the
Pelycosaurs still mainly within the nostril. In the Dinocephalians
(Delphinognathus, Tapinocephalus) it comes partly on to the face.
In the Dromasaurians (Galepus), it forms a very appreciable
portion of the facial wall; and in the Therocephalians (Seylaco-
saurus, Aloposaurus), it also appears pretty largely on the face.
In the Anomodonts it is absent, probably because they had lost
their organ of Jacobson, as would appear from the loss of the
prevomer.

In the Cynodonts, the septomaxillary is always present. In
the lower types it appears on the face, but in the higher forms it
is almost entirely inside the nostril.

Among Mammals a septomaxillary is known only in some of
the lower forms. In Yvitylodon it appears on the face between
the nasal and premaxillary, in much the same way as in Vytho-
saurus. In Ornithorhynchus and Echidna it would appear from
the researches of Gaupp that what used to be regarded as the
upper part of the premaxillary is really the septomaxillary. If
this be so, as seems pretty certain, then the Monotremes have the
septomaxillaries better developed than in the Cynodonts.

The only higher mammal in which there is a bone to be regarded
as probably the septomaxillary is Dasypus. Here a small bone,
which I described in 1897 asthe “ nasal-floor bone ” and suggested
might be homologous with the upper part of the premaxillary in
the Monotreme, is probably to be regarded as a rudimentary septo-
maxillary.

Vomer and Prevomers.

In 1895, and more fully in 1902, I showed that there was
reason to believe that the so-called reptilian ‘“‘ vomers” were not
homologous with the mammalian vomer, but that being formed as
splints to the paraseptal cartilages in close ae pceianion with the

63°

920 DR. R. BROOM ON THE

organs of Jacobson, they were really homologous with the bones
that unite to form the ‘‘dumb-bell bone” of Ornithorhynchus ;
and that the mammalian vomer had its homologue in the so-called
“‘narasphenoid”’ of the lower forms. As a new name was neces-
sary for the reptilian ‘‘ vomers,” I proposed the name prevomer.

In the Batrachia we find all three bones well developed, the
median true vomer or parasphenoid being especially large to
support the base of the skull. When in the earliest true Reptiles
the pterygoids came together, there was little need for the median
vomer and it became greatly reduced. In the Cotylosaurian
Diadectes the median vomer is still a fairly strong rod, but in the
later Cotylosaurians or primitive Diapsidans Pariotichus and
Procolophon the vomer is a very short pointed process. In most
later Diapsidans the vomer remains a small unimportant element.
It developes to a fair size in the Ophidia and becomes large and
much specialised in the Chelonia. The prevomers, on the other
hand, remain large in most Diapsidans, but where, as in the
Chelonians and Crocodilians, the organs of Jacobson become much
reduced or lost the prevomers likewise tend to disappear.

In the mammal-like Reptiles the vomer shows great variations.
In the Therocephalians it is small as in the primitive Diapsidans,
but with the development of a secondary palate a new function
is given to it, and it becomes large. In the Anomodonts, though
the secondary palate is only imperfectly formed, the vomer is large
and extends well forward. In the Cynodonts, where the secondary
palate is complete, the vomer is very large and extends from the
basisphenoid to near the front of the snout. The front part of
the bone corresponds so exactly in its relations to the mammalian
vomer, that it is impossible to doubt that the bones are homologous.
On the other hand, if the anterior part of the bone were lost it is
probable that every one would agree, from the relations of the
back part, that it was the homologue of the reptilian so-called
“‘parasphenoid.” In Mammals the vomer varies greatly in size.
It is relatively very large in the Cetacea, sometimes extending
from the basioccipital to the front of the rostrum, while in the
Rodentia it is often more or less rudimentary.

The prevomers are large in the Dinocephalians and Thero-
cephalians. In the Anomodonts they have completely disappeared.
In the Cynodonts, with the formation of the secondary palate
they are either greatly reduced (Gomphognathus) or quite absent
(Bauria). In Mammals the prevomers are usually absent, their
function as supports to Jacobson’s cartilages being taken by the
palatine processes of the premaxille. Im only two mammals are
they known for certain to occur as distinct bones, viz. Oriitho-
rhynchus and Miniopterus, and in both of these the pair of bones”
fuse together to form a median bone before the animal is full-
grown.

Some observations have recently been made which at first sight
appear to cast a little doubt on the homology of the parasphenoid
with the mammalian vomer.

SKULLS OF CYNODONT REPTILES. 921

Versluys has discovered what he believes to be a large para-
sphenoid in Dermochelys in addition to the vomer and in no way
connected with it. If this determination be correct, it will
probably turn out that the Chelonian vomer is after all a pair of
prevomers fused. The early development of the Chelonian vomer
has not, so far as I am aware, ever been examined, and in my paper
on the reptilian and mammalian vomerine bones I spoke very
guardedly on the subject. So far as we know, the Chelonian
vomer is always a median unpaired bone. But if it be a true
vomer, what of Versluys’ supposed parasphenoid? Fuchs has
shown that in Chelone the basisphenoid is ossified by a large
irregular exostosis on its under side, and that this exostosis bears
relations to the pterygoids very similar to those which the para-
sphenoid of Versluys does. In the light of the observations of
Fuchs, I think it must be concluded that the supposed parasphenoid
in Dermochelys is entirely a development of the basisphenoid, and
not the homologue of the parasphenoid of other reptiles.

Gaupp and Fuchs have both apparently discovered a rudimentary
ossification behind the vomer in Chelonians which they believe
to be a true parasphenoid, and Fuchs has discovered what he
believes to be a rudimentary parasphenoid in Didelphys. The
situation of these rudimentary ossifications is undoubtedly that of
the parasphenoid, but they are also in the region normally occupied
by the vomer in Mammals. When a bone which occupies one region
in an ancestor comes to occupy a somewhat different region in
a descendant through a portion of the bone becoming aborted, it
is by no means uncommon that rudimentary ossifications can be
detected in the region abandoned. The os caruncule-is undoubt-
edly the internasal process of the premaxilla in Ornithorhynchus
and Hehidna, but though it is quite detached from the pre-
maxilla, it is nevertheless a portion of the premaxilla. In the
case of the vomer, supernumerary ossifications appear to be not
uncommon both in front and behind. In Orycteropus there are
two small ossifications in front, apparently not prevomers, but
detached ossifications of the true vomer. Kitchen Parker seems
to have found them so commonly present that in some groups
he regarded them as the rule. Speaking of the condition in
Marsupials he says: “‘The main vomer is often relatively small ;
there is, nearly always, a pair of antero-lateral vomers....and
large postero-lateral, and other, or postero-medial vomers ; these
are very irregular and unsymmetrical in the young Cuscus
especially, in which I find ¢en vomerine bones.” Parker’s postero-
medial vomers are probably the ossifications regarded by Fuchs as
parasphenoids, and there seems no reason to regard them as of
any more morphological significance than the Wormian bones in
the human skull.

Alisphenoid.

Until recently the alisphenoid bone has been looked upon,
like the orbito-sphenoid, as an ossification of the cranial wall, and

922 DR. R. BROOM ON THE

according to Parker “ the alisphenoids and orbito-sphenoids appear
as chondrifications of the walls of the skull.” In studying the
development of the marsupial skull some years ago, I found that
the alisphenoid has originally nothing to do with the walls of the
skull. It first appears asa short rounded rod lying outside the
trabecula and quite independent of it or of any other skeletal
structure. In its relations it seems exactly to correspond to the
middle part of the cartilaginous bar on which the pterygoid bone
develops in Lizards and Sphenodon. In the majority of Lizards
this middle part gives rise to the epipterygoid or columella cranii.
When, asin Chameleon, the epipterygoid is rudimentary, the short
bar which forms its base is almost exactly similar in structure and
relations to the bar from which the alisphenoid developes in
Mammals. One therefore seems driven to the conclusion that the
epipterygoid and the alisphenoid are different developments of
the same element. And this conclusion seems borne out by
comparative anatomy, for we find that most Reptiles have either an
epipterygoid or an alisphenoid, but never both. In Lizards we
find an epipterygoid, but never an alisphenoid: in Snakes an
alisphenoid, but never an epipterygoid.

There seems little doubt that the epipterygoid is the early type
of development. We find, for example, in the primitive Proco-
lophon a columella cranii almost exactly like that of the lizard.
In the Therocephalians the columelle are long and slender, but
usually flattened. In the Anomodonts they are slender, but
rounded. In the Cynodonts we find no longer the columella
cranii, but in its place a broad fan-shaped alisphenoid. The
Cynodont alisphenoid further differs from the columella cranii
of the earlier forms in having the lower part well developed and
replacing the backward extension of the pterygoid. In the
mammal the alisphenoid differs from that of the Cynodont mainly
in having the 2nd and 3rd branches of nerve v. passing through
it instead of behind it.

[Vote by Kprror.—_In Dr. Broom’s memoir as presented to the
Society there followed here a discussion of the quadrate and
tympanic, illustrated by two diagrams. A recent discovery made
by Dr. Broom has considerably modified his views, and he has
asked leave to withdraw the paragraphs omitted here until he
has time to work out and present to the Society in a fuller
form the bearings of his new facts.— August 11th, 1911.]

Angular.

The angular is found in all mammal-like reptiles. It is large
in the Dinocephalians, Anomodonts, and Therocephalians, but
comparatively small in the Cynodonts. In Mammals there is a
small splint-bone on the lower side of Meckel’s cartilage which is
probably the remains of the angular. In Ornithorhynchus there
appear to be two splint-bones, one being probably the surangular.

SKULLS OF CYNODONT REPTILES. 923

In the following diagram is represented what appears to be the

genetic relationships of the principal known Cynodont genera :—

Therocephalian Ancestor.

Bauria

~~ Alurosuchus

“ie | ~G@alesaurus

=

Mammalhan xe
ape tor Ne
Ance stor. Sesamodon Nythosaurus

v Melinodon

Trirachodon

Cynognathus

SS
<
SS

=~ 4 va
Diademodon
Gomphognathus

List of References to the Principal Literature.

Baur, G.—On the Quadrate in the Mammalia. Q.J. Micr. Se. Vol. xxviii, 1886,
p. 169.

Broom, R.—On the Fate of the Quadrate in Mammals. Ann. Mag. N. Hist., Nov.
1890, p. 409.

— On the Occurrence of an apparent distinct Prevomer in Gomphognathus.
Journ. Anat. & Phys., Vol. xxxi, 1897, p. 277.

— On the Structure and Affinities of Udenodon. P.Z.S. 1901, Vol. ii,
p. 162.

— On the Classification of the Theriodontsand their Allies. Rep. 8. Afr. Assoc.
Ady. Se. 1903, p. 286.

—— On the Lower Jaw of a small Mammal from the Karroo Beds of Aliwal
North, South Africa. Geol. Mag., Aug. 19038, p. 345.

— On the Mammalian and Reptilian Vomerine Bones. Proc. Linn. Soc.
N.S. W., 1902, p. 545.

— Onthe Theriodonts in the Albany Museum. Rec. Alb. Mus., Vol. i, 1904, p.82.

— On the Structure of the Theriodont Mandible and on its Mode of Articu-
lation with the Skull. P.Z.S., 1904, Vol. i, p. 90.

— Preliminary Notice of some new Fossil Reptiles collected by Mr. Alfred
Brown at Aliwal North, S. Africa. Rec. Alb. Mus., Vol. i, 1905, p. 269.

(Preliminary notice of Sesamodon browni and Melinodon simus.)

924 DR. R. BROOM ON THE

Broom, R.—Reptiles of the Karroo Formation. In Rogers’ ‘ An Introd. to the
Geol. of Cape Colony.’ London, 1905, p. 228.

On some Points in the Anatomy of the Theriodont Reptile Diademodon.
P. ZS. 1905, Vol. i, p. 96.

—— Ona New Cynodont Reptile (42lurosuchus browni). Tr.S. Afr. Phil. Soc.,
Vol. xvi, p. 376.

—— Some recent Advances in South African Paleontology. Science, Vol. xxvi,
1907, p. 796.

— On the Origin of Mammals. Rep. Brit. & S. Af. Assoc. Adv. Sce., Vol. 1,
Johannesburg, 1907.

—— The Origin of the Mammal-like Reptiles. P.Z.S. 1907, p. 1047.

—— Observations on the Development of the Marsupial Skull. Proc. Linn. Soc.
N.S. W., 1909, p. 195.

—— Notice of some new South African Fossil Amphibians and Reptiles. Ann.

S. Afr. Mus., Vol. vii, 1909, p. 251.
(Description of Bawria cynops.)
A Comparison of the Permian Reptiles of North America with those of
South Africa. Bull. Am. Mus. Nat. Hist., Vol. xxviii, Art. 20, 1910, p. 197.

Doxto, L.—On the Malleus of the Lacertilia, and the Malar and Quadrate Bones
of Mammalia. Q.J.Micr. Sc. Vol. xxiii, 1883, p. 579.

Fucus, H.—Untersuchungen iiber die Entwicklung der Gehérknéchelchen, des
Squamosums und des Niefergelenkes der Siiugetiere, &c. Archiv f.
Anat. u. Phys., Anat. Abth., 1906.

—— Ueber einen Rest des Parasphenoids bei einem rezenten Siugetiere. Anat.
Anz., Bd. xxxii, 1908, p. 584.

-——— bBetrachtungen iiber die Schlafengegend am Schidel der Quadrupeda. Anat.
Anz., Bd. xxxv, 1909, p. 113.

— Ueber Knorpelbildung in Deckknochen, nebst Untersuchungen und Betracht-
ungen tiber Gehorknéchelchen, Kiefer und Kietergelenk der Wirbeltiere.
Archiv f. Anat. u. Phys., Anat. Abt., 1907.

—— Ueber das Pterygoid, Palatinum und Parasphenoid der Quadrupeden, &c.
Anat. Anz., Bd. xxxvi, 1910, p. 33.

Gapow, H.—On the Modifications of the First and Second Visceral Arches, with
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__ Trans. (B) Vol. clxxix, 1888, p. 451.
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von Hehidna aculeata var. typica. Abdr. Semon’s Zool. Forschungsreisen
in Australien. Jena, 1908.
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Mammalian Orders: with a Discussion of the Origin of the Mammalia
and of the Problem of the Auditory Ossicles. Bull. Am. Mus. Nat.
Hist., Vol. xxvii, 1910, p. 105.
Kinesiey, J. S.—The Ossicula Auditus. Tufts College Studies, No. 6, 1900, p. 203.
The Origin of the Mammals. Science, n. s., Vol. xiv, 1901, p. 193.
Ksniipere, K.—Beitrage zur Entwickelungsgeschichte des Kiefergelenkes. Ge-
genbaur’s Morph. Jahrb., Bd. xxx, 1904, p. 159.
Lusoscu, W.—Ueber das Kietergelenk der Monotremen. Jenaische Zeitsch. f.
Naturw., Bd. xli, 1906, p. 549.
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Am. Phil. Soe., Vol. xxiv, 1887, pp. 109-111,

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Remarks at the Discussion on the Origin of Mammals. Proc. Internat.
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Bia

SKULLS OF CYNODONT REPTILES. 925

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——. idem IX. Sect. 1. On the Therosuchia. Phil. Trans., Vol. clxxxv (1894)

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— Idem. IX. Sect. 2. The Reputed Mammals from the Karroo Formation
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—. idem IX. Sect. 4. On the Gemphodontia. Phil. Trans., Vol. clxxxvi (1895)

5 Do Ue

—— Idem. IX. Sect. 5. On the Skeleton in New Cynodontia frem the Karroo
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— On the Dentition of the Palate in the South African Fossil Reptile Genus
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1910, p. 487.

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Woopwarp, A. S.—Outlines of Vertebrate Palaeontology for Students of Zoo-
logy. Cambridge, 1898.

Explanation of lettering in the Plate and Text-figures.

Ang. Angular; Art. Articular; Aud.gr. Auditory groove; B.O. Basioccipital ;
B.S. Basisphenoid; Dent. Dentary; EH.O. Exoccipital; Ew.st. Extrastapedial ;
Fr. Frontal; Ju. Jugal; I.P. Interparietal; Ia. Lachrymal; Mail. Malleus ;
Men. Meniscus; Mx. Maxilla; Na. Nasal; O.O. Opisthotic; Pa. Parietal ;
Pal, Palatine; Pmx. Premaxilla; Po.O. Postorbital; Pr... Prefrontal; Pt.
Pterygoid; P.Vo. Prevomer; Qu. Quadrate; S.Ang. Surangular; Sma. Septo-
maxillary ; S.O. Supraoccipital ; Sq. Squamosal; Tym. Tympanic; Vo. Vomer.

EXPLANATION OF PLATE XLVI.

. Side view of the cranial wall of Cynognathus berryi. Half nat. size.

. Occiput of Cynognathus berryi. Half nat. size. :

. Side view ot the skull of Melinodon simus. Nat.size. This represents the
type and only known specimen. It is so badly crushed that it is
impossible to restore it with any confidence. Six molars of the right side
are preserved, and the axes of two molar series make with each other an
angle of about 60°. The frontal region is certainly narrow, and as the teeth
are very similar to those of Sesamodon it is probable that the skull is also
somewhat similar, but probably the snout is shorter in Melinodon and the
skull relatively broader.

. The molar teeth of the lett side of Sesamodon browni as preserved. X 1s.

. Base of the skull of Sesamodon browni as preserved. X 15.

. Quadrate and part of the squamosal of Bawria cynops, as seen from the
front. Nearly twice nat. size.

. Base of skull of Bawria cynops, viewed partly from the side to show the
deep keel of the basisphenoid.

. Occiput and base of skull of Bawria cynops, viewed from behind and partly
from below. Slightly restored. About two-thirds nat. size.

. Median section of skull of Diademodon. All parts in unbroken line are
from the specimen in the South African Museum. The prevomer and septo-
maxillary as restored are founded partly on the British Museum specimen
ot Gomphognathus and are partly hypothetical, the inner part of neither
bone bemg known in any specimen. About half nat. size.

gg

(SU Cl

Wey (2) NN Qoe

926 DR. A. HOPEWELL-SMITH AND DR. H. W. MARETT TIMS ON

39. Tooth-germs in the Wallaby AMacropus billardier.
By A. Hoprwett-Smirn, L.R.C.P., M.R.C.S., and
H. W. Marerr Tims, M.A., M.D., F.1.8., F.Z.S.,
King’s College, Cambridge.
[Received and Read May 23, 1911. |
(Plate XLVII.* and Text-figures 181-189.)

The dentition of the Marsupials offers, in some respects, points
of greater interest than does that of other mammals. Though
much has been already written upon this subject, there are still
problems awaiting solution.

The complete history of the tooth-genesis has been worked out
in few members of this class, due no doubt to the difficulty of
obtaining a sufficiently complete series of embryos of any one
species. Any contribution, therefore, however small, may become
of importance as forming a link in completing the chain of
evidence.

The material upon which these observations are based consisted
of three embryos of Macropus billardieri, obtained from the Seven
Sisters Islands by Mr. Brooke Nicholls of Melbourne, to whom
we are greatly indebted for his kindness in sending them for
examination. These embryos give the following measurements :—

Space | STace | Space
1 oe ee pase TI.
mm. mm. mm.
Tip of snout to occiput (circumferentially) ............) 21 | 85 38
3 35 5 (Gnistrare nteline) eeeeneseeeee 15 27 27
Tip of snout to root of tail (circumferentially) .........) 64 104 110
Vertex of head _,, 3 (amstraielit line) =... 35 | 66 52
CUESTA Ot tack not oly Mian HEN Sabai Rae blee Sabie he Alan THe: kp BL Dif
Total len¢th from tip of snout to tip of tail (cireum- |
ferential.) cetcaa cee aetna cae eaia eect nian 135 137

Serial sections were cut in a vertical transverse direction
though the skulls of Stages I and II and stained with borax
carmine. The jaws of Stage III, which gave measurements
approximately equal to those of Stage IT and presumably therefore
of an embryo of about the same age, were dissected and clarified
in oil of cloves after the method recommended by Huxley, to
show the relative positions of the calcified teeth.

The most detailed accounts of the development of the teeth in
Diprotodont Marsupials are those given by M. F. Woodward (12)

* For explanation of the Plate see p. 942.

Im, GS, US, Il, DAHVAUL.

TOOTH- GERMS IN MACROPUS BILLARDIERI.

TOOTH-GERMS IN A WALLABY. 927

and by Deppendorf (1); the latter, however, does not appear
to have examined any specimen of MJacropus. Woodward has
furnished the results of his examination of 14 embryos belonging
to four species, viz. I. giganteus, M. eugenit, M. bennetti, and
M. brachyurus. From the measurements given by him, allowing
for the differences in size of the adult animals of the different
species, our Stage I would appear to be considerably younger than
any of the embryos which he examined. Before discussing the
general considerations arising from our observations, we will
proceed to a description of the conditions found in our material.

Srace I.

Upper Jaw.—At the anterior extremity of the upper jaw the
oral epithelium sinks into the substance of the jaw, forming
a broad triangular cellular mass the outline of which is very
irregular. There are projections of some size extending laterally
into the premaxillary region. None of these would, by themselves,
suggest tooth vestiges, but the possibility of their being of such a
nature is not entir ely ‘negatived because, as will be shown in the
sequel, similar but more definite structures are to be seen in the
lower jaw. The evidence here is, however, too problematical to
permit of definite conclusions being drawn.

Text-fig. 181.

@
> e e \
e 2 * @ee ‘ n Nee
© \ an
89 oS z
ana 733 s0% Fy | ee a q =

ee ROR ee oe
= — Ge ~ = aoe
een OY =~ Neg i) me
Capaty Sen 9*00epccecgege oso
= = ~ = os = = =—_— «<>
Be eee een or!
ie ae oy ee ie
eee fi dew ey) ee Oral
i —~ eee sO 2 -
Fea ee ee arene = epithelium
ee a See heme pens ae
55 =aa

Section showing the tooth-band (X) passing horizontally inwards and forming
the anterior part of the tooth-germ B.

On either side of the middle line close to the nasal septum is a
minute slightly calcified tooth-germ with a dental papilla of

928 DR. A. HOPEWELL-SMITH AND DR. H. W. MARETT TIMS ON

definite shape. This germ will for the present be indicated by
the letter A.

From a point just external to the point of connection of the
neck of the enamel organ of 4 to the oral epithelium, the tooth-
band passes horizontally inwards towards the middle line
(text-fig. 181)*. This rapidly increases in size, forming on section
a conical mass of cells which soon loses all connection with the
oral epithelium. From the histological characters of the cells
and from the fact that in the more posterior sections a portion
of the stellate reticulum of the enamel organ is visible, this
structure must be regarded as a second tooth-germ (£).

It is considerably larger than the preceding germ 4, though
there is not the same amount of differentiation into the more
typical dental tissues.

Text-fig. 182.

eo

Section through the tooth-germ B at its deeper end.

The independent connection of the enamel organ 4 with the
oral epithelium shows that it is not to be regarded as the
morphological successor to A.

To neither A nor Bare there either palatal or labial down-

* Owing to the difficulty of accurate orientation, the anterior sections of a jaw cut
in the vertical-transverse direction will be in a plane parallel with the outer surface
of the jaw. This has been borne in mind in describing the relative positions of the
various structures.

TOOTH-GERMS IN A WALLABY. 929

growths which could serve as an indication as to which dentition
the germs should be referred.

As this tooth is traced backwards it assumes a peculiar shape on
section. Its inner margin is sharp, the outer one rounded, while
on the dorsal surface is a relatively deep fissure (text-fig. 182).

The peculiarity of the adult tooth is thus early indicated.

As B disappears from the sections a new tooth-germ (C) makes
its appearance. Though minute it has a fully formed enamel
organ and dental papilla, but without any trace of calcification.
This germ has a very superficial position in the jaw.

Closely following upon Cis a large uncalcified tooth (D) lying
to the palatal aspect of and slightly posterior to the preceding
tooth-germ. Connected with the neck of the enamel organ is a
minute predecessor slightly invaginated by a rudimentary dental
papilla (text-fig. 183).

Text-fig. 183.

<aye 088 One
Wa Noe x
AT

aay aye,

Section showing the tooth-germ D with its predecessor lying to the labial side
of the neck-band.

Yet another germ (/’) of some size soon makes its appearance,
occupying a superficial position in the jaw. There is a slight
bulging of. the dental lamina to the labial side of the neck of this
enamel organ, which may or may not indicate an abortive attempt
at the formation of a predecessor.

The dental lamina from this point is continued backwards for
some distance as a distinct band without showing any definite

930 DR. A. HOPEWELL-SMITH AND DR. H. W. MARETY TIMS ON

trace of a tooth-germ. After an interval, another tooth (//) of
considerable size appears, lasting through three slides. It has
a dental papilla with blunted apex. As yet there is no definite
evidence of calcification. There are neither palatal nor labial
downgrowths of the dental lamina. ‘This tooth we identify as
the first of the cheek series.

The tooth-band continues backwards in a well-marked condition
giving evidence of a distinct swelling which in Stage II has
developed into an undoubted tooth-germ. Without a knowledge
of the subsequent history of this swelling, one would have
hesitated to attach importance to it.

Posteriorly to this is the large enamel organ of Stage I from the
neck of which springs a definite palatal downgrowth of the dental
lamina. The shape of the dental papilla is definitely molariform,
the outer cusp already attaining to a higher level than the inner.

One more enamel organ (/), the most posterior in the jaw
at this stage, is to be seen following closely upon the preceding
one. It has but reached the flask-shaped stage.

Thus it is seen that there are representatives of eight teeth in the
upper jaw at this stage which, by comparison with those present
in Stage II, we identify as incisors 1, 2, 3, 4 and 6, and premolars
1,3 and4. The reasons which have led us to arrive at this con-
clusion will be detailed subsequently.

Lower Jaw.—Near to the mandibular symphysis is a well-
defined involution of the oral epithelium into the subjacent
mesoblastic tissues. The cells, both peripheral and central,
of this tubular downgrowth have precisely the same characters
as those covering the alveolar margin. From the appearance of
these cells and from the inclination of the downgrowth (down-
wards and towards the middle line) being the same as that of
undoubted tooth-germs situated more posteriorly in the jaw, the
opinion that it is a vestigial tooth-germ (a) seems to be justified.
Very closely following upon this structure and placed at a deeper
level in the jaw, is a small well-calcified tooth (6) showing both
dentine and enamel. The appearance of this tooth suggests
degeneration, and it may safely be asserted that this tooth, though
calcified, is not the one which will ultimately become the functional
incisor.

The difficulty in identifying the exact relationships of tooth-
germs close to the symphysis is always great, due in part to a
certain amount of crowding, accentuated in the Macropodide by
the very large size of the functional incisor and in part to the
difficulty of correct orientation. We at first thought that these
two structures stood to each other in the morphological relation-
ship of predecessor and successor. From a careful consideration
of all the facts we believe that this is not the case, but that they
are the representatives of two separate teeth and that the close
approximation to each other is due to the crowding caused by the
large functional incisor. The appearance of @ suggests a tooth

TOOTH-GERMS IN A WALLABY. 931

which is attempting but failing to develope; it has not the
“concentric ” appearance of a vestigial tooth-germ such as one
so often sees as the last evidence of existence of a predecessor to a
calcified tooth.

Rapidly succeeding the calcified tooth 6 is a well-defined flask-
shaped dental rudiment lying in a very superficial position in the
jaw. This, though close to, is quite independent of any of the
other tooth-germs here present and will in the meanwhile be
referred to by the letter c (text-fig. 184).

Text-fig. 184.

- oe
° 7 OC = He
— Ze. mae 4 2 ose,
Sue Wig > oe oyguant™
23TH 2 @ o%7 ye 0)
= 12 2,2 «© = Ww
as — » 4,009 9a68° ¢
sis ea . ?
= >= POS GSS °
1 oF G wes, 1 - ,
1 TNT GMRRR SS
— ? Le pe O &., 0-7
oe 5 7s eee, 9 0
roy ’ aay, er. - yen
r} eee 1 00 te Des! NDbibin D
A eo? oe 0 ’ Be Ps
’ 6 .
ast, ovoy. ; 0%
ry

Section showing flask-shaped rudiment of tooth-germ c.

At this point the dental lamina runs horizontally inwards, as
in the upper jaw. It forms a thickened band of cells lying some
distance below the tooth-germ c. Tracing this band backwards
it quickly becomes connected with the upper part of the enamel
organ of a large tooth (d) in which the dental tissues are differen-
tiated and which extends through several slides. In relation to
the neck of this enamel organ is a lingually situated downgrowth
of the dental lamina indicating a potential successor (ds) to the
tooth d. From this poimt the tooth-band vanishes for some
distance (text-fig. 185).

As the tooth itself dies out the dental lamina once more comes
into prominence and gives rise to an enlargement indicating in our

DR. A. HOPEWELL-SMITH AND DR. H. W. MARETY TIMS ON

932

Text-fig. 185.

end of tooth-germ ¢ and commencement of enamel

Section through posterior

an of d and rudiment of a successor to d.

org

Text-fig. 186.

Section showing the enlarged downgrowth of the dental lamina indicating the

vestige of the tooth-germ e.

TOOTH-GERMS IN A WALLABY. 933

opinion a tooth vestige (e). It lies close beneath the alveolar
margin (text-fig. 186).

Then occurs a further interval through which the tooth-band
persists without giving rise to any further dental enlargements.
Three further teeth (h, 7, and 7) of premolariform pattern, follow
in the cheek region. Of these the middle one is considerably
the most extensive, the posterior one being quite small. To the
two anterior teeth there are indefinite indications of lingual
downgrowths of the dental lamina.

As in the upper jaw there are indications of eight teeth, five
antemolars and three maxillary teeth,

A general impression of the number and relative positions of
the dental structures in both jaws may perhaps be best realised
by the following diagram, which has been drawn to scale in the
horizontal direction.

Text-fig. 187.

Diagram showing number and relative positions of the tooth-germs in the jaws
of Stage I. Calcified germs more heavily outlined.

Srace IT.

The difference in size between this embryo and that of Stage I
is very considerable, and as a consequence a number of additional
teeth have appeared in both Jaws and those which were present
in the earlier stage have developed very materially. It is un-
fortunate that we have not had at our disposal an embryo of

Proc. Zoou. Soc.—1911, No. LXIV. 64

934 DR. A. HOPEWELL-SMITH AND DR. H. W. MARETT TIMS ON

intermediate size, as this would have rendered the precise
interpretation a matter of less difficulty and of greater certainty.

Without going into details we will summarise as briefly as
possible the conditions found in the later stage.

Upper Jaw.—Anteriorly is a small calcified tooth (4’) oceupying
a position similar to the tooth found in the previous stage and is
undoubtedly the same tooth still persisting. It shows no signs of
being erupted, neither has it undergone any further development.
That it will ultimately become absorbed without attaining fune-
tional activity seems to be certain.

Behind this hes a large incisiform tooth already heavily
calcified. This tooth (b') is the largest of the anterior series.
All traces of the tooth-band between dA’ and 45’ have become
lost, and several sections intervene between the disappearance
of A’ and the commencement of the enamel organ of B. Thus
we are of the decided opinion that these are two morphologically
distinct teeth, a point to which reference will be made later.
This conclusion is the same as that at which we arrived in Stage I.

C" is still a small unealeified tooth lying superficially, and
appears to have been pushed out of the serial line towards the
outer aspect of the jaw.

D' is very similar to 5, but perhaps not quite so large.
Posteriorly to )’ is a small calcified tooth (4’) of irregular shape
and situated close to the alveolar margin of the jaw. ‘This tooth
is obviously vestigial and will never become functionally active.
According to our identification this tooth is unrepresented in the
earlier stage.

After an interval another large incisiform tooth is seen (2”).
It is neither so large nor so heavily calcified as B’ or D’.

G' is a tooth of some size showing but traces of commencing
calcification. From its relation to the premaxillo-maxillary
suture we regard this tooth as the canine. It is a tooth in an
early stage of development which has appeared since the age of
Stage I. There is a ‘“‘ concentric epithelial body” which is to be
regarded as the last trace of a vestigial predecessor.

The first of the true maxillary teeth is elongated and of
premolariform type (H’) without any indications of a predecessor
or successor. The third tooth (/’) is like unto it but of much
larger size, extending through many more sections.

Between these two calcified teeth is a very deeply placed tooth-
germ with a neck of remarkable length (/)) (Pl. XLVII. fig. 1).
‘here is no evidence of calcification. This is evidently the
second maxillary tooth which, crowded between the large calcified
first and third, has been forced into the abnormally deep situation,
besides being pushed from the linear series towards the palatal
side of the jaw.

The fourth maxillary tooth (A’) is large and in it calcification
is but just commencing. From the neck of the enamel organ
springs a marked labial downgrowth of the dental lamina, con-
nected with which is a ‘concentric epithelial body,” the vestige

TOOTH-GERMS IN A WALLABY. 935

of a predecessor (Pl. XLVIT. figs. 2. & 3). ‘The importance of
this will be discussed later. The last tooth of the series (Z’) is
uncalcified and at an earlier stage of development than the
maxillary teeth in front of it. It lies much nearer to the alveolar
margin than do the others, and there is this further point of
interest, that the anterior end of this tooth overlaps superficially
the posterior end of the tooth in front, which is distinctly not the
case with the more anteriorly situated teeth.

According to our identification we recognise at this stage six
incisors, one canine and five maxillary teeth.

Lower Jaw.—The difficulty of interpreting the appearances in the
region of the mandibular symphysis is even greater than in the
earlier stage. The enormous development of the lower functional
incisor, extending as it does through thirty slides*, has disarranged
all the parts, more particularly anteriorly where the tip of the
tooth becomes more superficial and where it now reaches well in
front of the point at which its young enamel organ connected with
the oral epithelium.

Text-fig. 188.

Section showing involution of the oral epithelium
at the mandibular symphysis.

At the symphysis the oral epithelium is turned inwards in the
middle line so as to partially cover the opposing ends of the
mandibular rami. At this point there is a definite involution
of the epithelium into the subjacent tissues. The section being
cut at this point in a plane parallel with the anterior surface, the

* A rough estimate may be obtained by stating that a slide carries an average
of twelve sections, each section being 745 mm. in thickness.
: 64*

936 DR. A. HOPEWELL-SMITH AND DR. H. W. MARETT TIMS ON

direction of this digitiform involution is inwards and _ slightly
upwards. ‘There is an involution on either side of the symphysis
and they are quite symmetrical. The position makes the inter-
pretation somewhat doubtful, otherwise we should have but little
doubt in ranking them as tooth rudiments. Regarding the origin,
connection, and histological characters as ef more importance
than mere position, we shall regard it as such and provisionally
indicate it by the letter a’ (text-fig. 188).

Close to the middle line, but on the alveolar margin of the
mandible, is a more globular involution containing cells commenc-
ing to be arranged as a concentrical epithelial body such as several
authors now recognise as being tooth vestiges. This will be
referred to as 6’. This also is a bilaterally symmetrical structure
and extends through several sections. Immediately above this is
a fissure in the oral epithelium containing some deeply stained
fibrous-looking material. The appearance suggests the possibility
of this marking the situation where the minute calcified tooth,
present in the jaw in this position 1m the earlier stage, may have
been erupted. There is otherwise no trace of the calcified vestige.

Text-fig. 189.

Showing the relative positions of the teeth in Stage IT.

Calcified teeth more heavily outlined.

A broad bulbous involution of the oral epithelium is seen close
to the middle line (c’) which soon loses its connection with the

TOOTH-GERMS IN A WALLABY. 937

alveolar epithelium, remaining as an isolated spherical mass.
It probably is the same structure described in Stage I, and is
naturally of larger size.

Lying ata deeper level but entirely independent of it, is the
anterior extremity of the large calcified functional incisor. That
these two structures have no morphological relationship with each
other is made clear, not only by their relative position but also
by the fact that the connection of the enamel organ with the
surface is definitely visible in a section farther back, together
with a “ concentric” vestige of a predecessor and an indication of
a potential successor. This large functional incisor we shall
indicate as d’,

Then follows an interval of thirteen slides without any trace of
teeth, and throughout the greater part of this interval the dental
lamina is scarcely visible. At length a definite “concentric
epithelial mass” occurs embedded in the dental lamina. It is im-
possible to say whether this is the vestige of a canine or of an outer
incisor. From its position relative to that of the opposing teeth, it
is probably the vestige of the latter tooth. The point in favour of
its being a vestigial canine is its propinquity to the first true post-
canine tooth. However, it is a point of quite minor importance.

The description already given of the true maxillary teeth
applies almost equally well to the corresponding structures in the
lower jaw. The only two points to which attention may be drawn
are (i) the larger size and definite calcification of the 4th tooth of
the lower series; and (ii) the absence of a definite ‘“ concentric”
predecessor to the same tooth, though there is a very minute
structure which may indicate its remains, the position in relation
to the tooth itself being the correct one. That it should have
disappeared is what would be expected from the greater size and
calcification of the successional tooth.

Jil, Hisronoey.

There are certain histological peculiarities revealed by an
examination of these jaws. They are briefly as follows :—

(i) The oral epithelium along the alveolar margins is very thick
and heaped up to an unusual extent. In no other mammal that
we have examined have we ever seen this character carried to such
excess. This is the ‘‘Zahnwall” of the German writers. It is
said to be a marked character in the Ungulates, and in our ex-
perience it is certainly not of common occurrence among manimalia
generally. Of course no morphological significance is implied
by the comparison.

(ii) The somewhat unusual compactness of the stellate reticulum

and the definiteness of the stratum intermedium of the enamel
organ.

(iii) The precocious developinent of the enamel which seems
to calcify as soon as the dentine, and even in some cases to

938 DR. A. HOPEWELL-SMITH AND DR. H. W. MARELT TIMS ON

precede it. Spherules of calco-globulin are clearly seen in many
of the ameloblasts.

(iv) The abundant evidence of blood-vessels within the enamel
organ is a very striking feature (Pl. XLVII. fig. 5). The fact, of
the vascularity of the enamel organ is one upon which there
are contradictory statements. Wedl, Magitot, Legros, Sudduth
and Paul deny the existence of blood-vessels in this situation.
On the other hand, Prof. Poulton and the late Prof. G. B. Howes
have mentioned their presence in the enamel organs of the Rat.

In Macropus there is no doubt whatever. They can be seen
entering the enamel organ apparently at more than one point on
the surface, and are often of sufficient size to clearly recognise the
blood corpuscles within them. They can be traced to a point about
midway between the outer and inner enamel epithelium, but we
have not seen them proceeding farther, neither have we found them
in the stratum intermedium, the position in which they were
described by the authors above mentioned.

The fact of the vascularity of the enamel organ is one of im-
portance in connection with the various theories held as to the
processes of the calcification and enamel formation and of the
functions of the stellate reticulum. It is extr emely probable that
the unusual vascularity of the enamel organs in this animal is
correlated with the precocious development of the enamel, to which
reference has just been made.

(v) Evidence of the fusion of enamel organs. There appears to
be some evidence of the possibility of the occurrence of such fusion
though the evidence is not decisive. Pl. XLVII. fig. 4, shows the
subdivision of an enamel organ into two parts by an epithelial
septum passing from the outer enamel epithelium to the inner
where the latter lies over the apex of the dentinal papilla. This
occurs in more than one cheek-tooth and may be seen on both sides.
We have never met with anything of the kind before. ‘That this
septum is not a small capillary running directly across the stellate
reticulum we are assured. Wecan offer no other suggestion than
that it is a double enamel organ taking part in the formation of a
single tooth.

This is a point of considerable morphological importance as
bearing upon the question of the evolution of the molar patterns.
Fusion is presupposed to have occurred by all writers who believe
in any form of concrescence, but little satistactory ev idence has
been forthcoming in the mammalia and little weight can be given
to the evidence derived from the disintegration ‘of the Cetacean
molars.

Amongst the lower vertebrates the evidences of fusion seem to
be conclusive. In one family of upper Paleozoic Sharks, the
Cochliodontide, there is a fusion into plates not only of adjoining
teeth of one series, but also of successional teeth of several series.
Semon (4) has also shown that a fusion of individual cusps takes
place in Ceratodus, and Rose has seen the cusps in the process of
fusion in the teeth of achameleon (3). Woodward (/oe, cit.) speaks

TOOTH-GERMS IN A WALLABY. 939

of asupernumerary incisor in the upper jaw of anadult Perameles
in the teaching collection of the Royal College of Science in which
there are “ indications of at least three fangs, and is obviously a
fusion of (these) teeth.” The possibility of tooth fusion in the
mammalia is therefore by no means improbable. The matiter
must remain unsettled until more satisfactory evidence is forth-
coming, but the condition figured would seem to point in that
direction and we can only repeat that we can suggest no other
explanation.

LV. GENERAL CONSIDERATIONS.

_ A. Serial homologies of the Teeth.

Upper Incisors.—Assuming our identification of the teeth to be
correct, we have six incisors present in the upper jaw, of which
the 5th is unrepresented in the earlier stage and has attained but
a slight degree of development in Stage IJ. This excessive
number of upper incisors confirms the original statement of M. F.
Woodward (11), as the result of his observations on Petrogale
penicillata and Macropus giganteus. That author is obviously
aware of the difficulty in accepting so large a number, for he wr ites
‘(p. 465): “The discovery of six pairs of incisors, although an abso-
lute fact, is in many respects an unfortunate one, as we know
of no adult mammal with so many, and even amongst Reptiles,
many Lizards and Crocodiles have the number of teeth in each
premaxilla restricted to five.” He then proceeds to discuss various
possible explanations such as the late calcification of the functional
incisors in Jacropus and the greater space in the premaxillary
region in Petrogale as the possible determining causes. A reference
to our text-figures 187 & 189 (pp. 933 & 936) will show that
Woodward has possibly found the correct explanation, both
factors appearing to be present in our specimens.

The difficulty is, however, not fully stated by a mere reference
to recent forms, for Osborn (2) in his paper, “On the Structure
and Classification of Mesozoic Mammalia,” considers that the
dental formula of the primitive heterodont mammal should be
taken as 1.4, c. 1, pm. 4, m. 8.

In a paper published three years later (13) Woodward appears
to have modified his former opinion, for in spite of having pre-
viously spoken of the presence of six upper incisors as ‘fan Alcolite
fact,” he writes “ Tam now disposed to regard the three minute
teeth which I described in the upper jaw of several genera
belonging to this family” (¢.e. Macropodide) “as the premilk
predecessors of the three functional incisors.” He gives no reasons
for the alteration in his opinion, as in his previous paper he dis-
cusses and discards the interpretation which he appears later to
have adopted. He further makes no reference to the alteration
in the identification of the homologies of the functional incisors
which such a change of opinion would involve.

We have naturally therefore paid particular attention to these

940 DR. A. HOPEWELL-SMITH AND DR. H. W. MARETT TIMS ON

‘views, and we agree with Woodward’s first statement that the
presence of six upper incisors, in spite of all the difficulties which
it involves, is ‘‘an absolute fact.” The one point upon which we
join issue with Woodward is his view of the homology of the first
functional incisor which he regards as being i. 1, whilst we hold
that it is i. 2. The small calcified vestige in both our stages
is clearly anterior to and independent of the first large tooth
which is obviously the first functional incisor. Our disagree-
ment is due to differences of interpretation of the conditions
present in a very puzzling region, 7%. e. close to the median
symphysis; and though we hesitate to put ourselves in opposition
to so accurate and experienced an observer, we nevertheless are
compelled to adhere to the opinions already stated. Weare, how-
ever, quite in accord with him in regarding the outer functional
incisors as being the 4th and 6th, the 3rd and 5th disappearing.

We, like Woodward, are therefore at variance with the opinion
of Oldfield Thomas (5), who regards the missing incisors as being
the outermost ones of the series. Though Deppendorf (1) does
not appear to have examined the jaw of MJacropus, he has made
interesting observations on the tooth-genesis in many other mar-
supials. In #pyprymnus he identifies the three functional incisors
as the 2nd, 3rd, and 5th. Thus he agrees with us in the identifi-
cation of the 1st functional teeth, but differs from both Woodward
and ourselves in the identification of the two outermost functional
premaxillary teeth. It is unfortunate that there should thus be
four different versions given on this subject. Possibly the con-
ditions described by Deppendorf may be really different and not
merely a difference in interpretation, as he carried out his
researches in other genera, in which the relative proportions of
the bone may be somewhat different and thus produce different
results.

Lower Incisors—In the lower jaw we believe there are evidences
of five ante-molar teeth. Whether the most posterior one is the
vestige of a canine or of an outer incisor, it is impossible to deter-
mine, Of theremaining four the first three are vestigial, so that
according to our interpretation the functional incisor of the adult

‘is i. 4. This statement is at variance with Woodward’s conclusion
as to the homology of this tooth which he regards as i.2. In his
illustration he figures two vestiges which obviously correspond
with our b' and ¢’, his first incisor in Petrogale being a minute
calcified tooth just as is the germ 6’. The morphological 3rd incisor
of Petrogale is “ more vestigial than the first,” just as c’ is more
vestigial than 6’.

The discrepancies between our interpretations probably depend
on the following facts. (1) Woodward seems to have examined the
jaw on one side only, and to do so would have divided it through at
the symphysis. In doing so he would most certainly destroy the
minute vestige a which we have found lying practicaily within
the symphysis. (2) He regards the posterior vestige as being
ete as to the large functional incisor, whereas we regard it as

eing anterior. We both agree in stating that it lies close to the

TOOTH-GERMS IN A WALLABY. 941

tip of the large tooth, and there is no doubt that the exact mor-
phological localisation of the vestige in relation to the large incisor
is a matter of difficulty. On the whole, therefore, we are in closer
agreement with Woodward's conclusions than with those of any
other writer.

The close approximation which we have found between the
representatives of these four anterior incisors is of considerable
interest when we remember the condition of the lower incisors in
Didelphys.

Canines.—These teeth call for no further observations.

Maxillary teeth—The fourth true maxillary tooth of the upper

jaw, as has already been stated, has a well-marked ‘ concentric
epithelial body” lying superficially to it. The importance of
these structures as representing the last trace of a tooth vestige
has been frequently insisted upon by one of us (H. W.M. T.)
(7,8), and the more recent researches of Wilson & Hill (10) seem
to have placed the matter beyond doubt. So long as the present
distinction as to whether or not there are predecessors is the cri-
terion of distinction between premolars and molars, confusion is
bound to result. Embryology often reveals the presence of these
vestiges which breaks down this conception of a true molar tooth.
It would in our opinion be better to call all post-canine teeth
maxillary teeth. Until such a suggestion is adopted we must
regard the fourth tooth of the series as a premolar (pm. 4).
The relationship of this tooth to ae one immediately behind it
suggests that the fifth tooth, 7. e. m’, is the deciduous tooth which
is replaced by pm’, a suggestion strongly urged by one of us in a
previous paper (8).
_ It seems to be tolerably certain that the 2nd maxillary tooth
will not be able to develop further, thus confirming the opinion of
Oldfield Thomas that pm? is the missing pr emolen,

The same interpretation holds good io the lower teeth.

To which dentition do the teeth of Marsupials belong ?

No very decided answer to this question can be obtained from
the material examined. ‘Traces of a successional dentition are
scanty, while those of a deciduous series are somewhat more
abundant and more distinct. One of us (H.W. M.'T.) has in
more than one paper (6, 7) previously urged, upon general grounds,
the view that the permanent dentition of the marsupials is the
permanent dentition of the Eutheria and that both the deciduous
and successional pm* belong to one and the same dentition. This
conclusion has been independently arrived at by Wilson & Hill
(10) and adopted by C. 8. Tomes (9). The joint authors of the
present paper are in agreement upon this subject, and we have
found nothing in the material which we have examined to cause
us to doubt the correctness of this view.

Note.-—I would desire to express my gratitude to the Odonto-
logical Society of Great Britain, for a grant in aid of researches

942

ON TOOTH-GERMS IN A WALLABY.

upon mammalian dentitions, and which has in part been used for
the purposes of this investigation ——H. W. M. '.

LITERATURE.

1. Deprenporr, T.—‘ Zur Entwickelungsgeschichte des Zahn-

3.

1

12.
13.

Fig.
Fig.
Fig.
Fig.
Fig.

systems fete Mar supialier.” Semon’s Zoolog. For schungs-
reisen in Australien und dem malayischen Archipel,” Bd. ill.

1897-1910, pp. 243-402.

. Osporn, H. F.—‘ On the Structure and Classification of the

Mesozoic Mammalia.” Journ. Acad. Nat. Sci. Philadelphia,
vol. ix. p. 186.

Ross, C.— Ueber die Zahnentwickelung vom Chameleon.”
ia Anzeiger, Bd. vil. 1893, pp. 566— B77.

. Semon, R.—‘Die tiussere Hingwiekelun g des Ceratodus forsteri.”

Zoolog. Forschungsreisen in Australien und dem malayischen
Archipel, TBials te 1893.

THomAs, OLDFIELD.—‘‘ On the homologies and succession of
the Teeth in the Dasyuride, ete.” Phil. Trans. Lond.,
vol. 178, 1888, pp. 443-462.

Rint e Ww. Lassa, — Notes on the Dentition of the Dog.”

eae Anzeiger, Bd. xi, 1896, pp. 537-546.

“The evolution of the Teeth inthe Mammalia.
Anat. & Phys. vol. xxxvii. 1903, pp. 132-149.

“On the succession and homologies of the Molar and
Premolar Teeth in the Mammalia.” Jour. Anat. & Phys.
vol. xxxvi, 1902, pp. 321-343.

th)

Journ.

. Tomes, C. S.—“ Dental Anatomy.” London, 1898.
. Witson, J. T., & Hitt, J. P.—‘“ Observations upon the

development and succession of the Teeth in Perameles,
together with a Contribution to the discussion of the homo:
logies of the teeth in Marsupial animals.” Quart. Journ.
Mier. Sci. Lond., vol. xxxix. n.s., 1896-97, pp. 427-584.

“ Observations on tooth development in Orni-
thorhynchus.” Quart. Journ. Micr. Sei. Lond., vol. li. n.s.,
1907, pp. 137-165.

Woopwarp, M. F.—‘‘ On the development of the Teeth in the
Macropodide.” Proc. Zool. Soc. Lond. 1893, pp. 450-471.
“On the Teeth of the Marsupialia, with especial
reference to the pre-milk dentition.” Anat. Anzeiger,
Bd. xii. 1896, pp. 281-291.

EXPLANATION OF PLATE XLVII.

1. Showing the deeply situated bulbous rudiment of pm? with the very long
neck of the enamel organ reaching to the surface of the oral epithelium.

2. The enamel organ of pm with the labial dow ngrowth of the dental lamina
near to the free extremity of which is an enlargement, a “concentric epi-
thelial body ” (c.e.b.).

8. A ‘‘ concentric epithelial body ” more highly magnified.

4, Showing the sub-division of the enamel organ into two parts—?a fusion of
two distinct enamel organs.

5. Showing the presence of bloodevessels w aie the stellate reticulum of an
enamel organ.

P.Z.9.1944. Pl. XLVILL.

G.M.Woodward delet lith. West, Newman imp-

IDION(O BISA EMCI IalO) lee G37

DR. C. W. ANDREWS ON A NEW FOSSIL MAMMAL, 943

40, On a New Species of Dinotherium (Dinotherium hobley:™)
from British Hast Africa. By C. W. AnprEws, D.NSc.,
F.R.S., F.Z.8. (British Museum, Natural History) f.

[Received & Read May 23, 1911.]
(Plate XLVIIT.F)

During the last few years great additions have been made to
our knowledge of the extinct mammalia of Africa, but hitherto
the discoveries of their remains have been confined to the
northern and southern portions of the continent. Now, however,
a new find of mammalian bones in British Hast Africa shows
that great hopes may be entertained that before long light will
be thrown on the history of the group in the central portions of
the continent. Recently Mr. C. W. Hobley, C.M.G., Commissioner
of Mines in British East Africa, sent to the British Museum a
small box of bones from the neighbourhood of Karungu on the
east side of Lake Victoria Nyanza. Most of the specimens are
indeterminable fragments, probably picked up on the surface,
but in addition to these there are some beautifully preserved
teeth with a portion of the mandible (Pl. XLVIII. figs. 1, 1 a) of
a small species of Dinotherium; a small imperfect Proboscidean
caleaneum (fig, 5), a patella, and some other fragments probably
belong to the same animal. The teeth and bone are in a wonder-
fully good state of preservation, being hard and not easily broken,
and there can be no doubt that further collecting in the same
locality will yield results of the very highest importance and
‘interest. '

The teeth preserved all belong to the lower jaw: they are
pm. 4, m. 2, and m. 3 of the left side, and pm. 3 of the right; the
left pm. 4 and m. 2 have been replaced in their sockets in the
portion of the jaw preserved; this also contains the freshly
broken roots of pm. 3 and m. 13 m. 3 is isolated, the portion of
the jaw behind m. 2 being lost. Anteriorly the jaw fragment ceases
-at the level of the front of pm. 2, where its inner border is turning
in towards the symphysis; its ventral portion is missing, the
dental canal being exposed.

The teeth.—'The anterior premolar (figs. 3, 4) has a crown con-
sisting of a high, laterally compressed antero-external cusp, to the
inner face of which a shorter and more rounded antero-internal
cusp is closely united. The hinder half of the tooth consists of a
comparatively low and compressed outer tubercle separated from
‘the antero-external cusp by a notch, and a small rounded inner

* [The complete account of this new species appears here, but as the name and
a preliminary diagnosis were published in the ‘ Abstract’ the species is distinguished
by the name being under'lined.—Eprror. |

+ Published by permission of the Trustees of the British Museum.

t For explanation of the Plate sce p. 945,

944 DR. C. W. ANDREWS ON A NEW FOSSIL MAMMAL,

tubercle which is joined to the outer by a ridge widening from
within outwards,

The last premolar (pm. 4) (figs. 1, 1 @) consists essentially of two
transverse crests which are united towards the outer side by a low
obscure ridge. On the anterior face of the tooth there is a slight
prominence of the cingulum, the surface of wear of which becomes
continuous with that of the outer end of the anterior crest.
There is also a low transverse ridge formed by the cingulum”
on the posterior face of the tooth.

The first molar has unfortunately been lost; the second (m. 2)
consists of a pair of transverse crests, the ends of which, especially
the inner end, are higher than the middle and the anterior face
of which is concave. Qn the posterior end there isa well-developed
ridge of the cingulum extending nearly the whole width of the
tooth. The third molar (m. 3) is similar to the second except
that it has a well-developed triangular talon, the outer side of
which is formed by a prominent tubercle from which a crest
diminishing in height and width runs to the inner edge of the
tooth,

In size and, on the whole, in the pattern of the teeth this
Dinotherium is very similar to ). cwviert Kaup, which is from the
lower and middle Miocene of France, being apparently especially
characteristic of the Burdigalien horizon. Detailed comparison
however shows some differences. In pm. 3 the separation of the
antero-internal cusp is much more distinct than in a specimen of
the same tooth ascribed to D. cwviert ; the last premolar is shorter
in proportion to its width than in that species, and in the talon of
m. 3 the outer tubercle is much more distinctly and independently
developed. These differences, coupled with the remoteness of the
localities in which the two forms are found, seem to justify the
establishment of a new species for this East African animal and |
propose that it shall be called

DINOTHERIUM HOBLEYI Andrews.

Abstract, P. Z.5. 1911, p. 35 (May 30th).

The dimensions (in centimetres) of the teeth in the type-
specimen are :—

Length. Width.
[DIU Se eee: 4-0 32
Ole Ape cance 4°9 4-2
U1 Ob, PAIRS rec 6:2 5:2
MUO ee ee 02 54

An imperfect proboscidean caleaneum (fig. 5) probably belongs
to this species. It belongs to the left side, and a great part of its
outer half is broken away and some of the other faces are abraded.
Its extreme length is 17:2 em., the cast of a caleaneum referred
to Dinotherium giganteum is 32°7 em. or nearly twice as long.

From the same locality there have been obtained a lower molar

ON RATS FROM THE SOCIETY’S GARDENS. 945

and the distal end of a radius of a small species of Rhinoceros
(? Aceratherium), portions of the carapace of a very large
Chelonian, probably a species of Zestudo, fragments of the shell
of Trionyx, and some Crocodilian remains.

From some portions of matrix adherent to some of the bones it
appears that they are preserved in a bed of tough clay with much
calcareous matter and numerous grains of blown sand ; this deposit
is probably of lacustrine origin, but in the absence of any molluscs
or other invertebrates, it is not possible to be certain either as to
its origin or as to its exact age. Judging from the Dinotheriwm
remains, the beds are probably lower or middle Miocene. If they
should fortunately turn out to contain a rich mammalian fauna,
probably this discovery will lead to a great advance in our
knowledge of the history of several groups of Artiodactyls, of
the Hyracoids, and possibly of the Anthropoidea. It is greatly
to be desired that a careful collection should be made as soon
as possible.

EXPLANATION OF PLATE XLVIII.

Fig.1. Dinotherium hableyi, portion of left ramus of mandible with pm. 4, m. 2, m. 3,
trom above; la, ditto, from side. Type specimen.

. Ditto, crown of m.2 of another individual.

. Ditto, pm. 3 of the type specimen trom above ; 3 a, ditto, from side.

Ditto, crown of pm.3 of another individual.

Ditto, left calecaneum from inner side.

Om oo to

eub., facet for cuboid ; ect., ectal facet ; sus., sustentacular facet ; é.c., tuber calcis.

All the figures are ? natural size.

EXHIBITIONS AND NOTICES.

June 13, 1911.
Epwin T. Newton, Esq., F.R.S., in the Chair.
Mr. H. G. Puimmer, F.RB.S., F.Z.S., Pathologist to the Society,

presented a Report on the Pathological Examination of Rats
(Mus decumanus) caught in the Regent’s Park and in the Society’s
Gardens. 500 rats had been examined between the Ist of J anuary
and the 17th of May, 1911, all in a precisely similar manner, The
spleen, lungs, glands, and blood were examined microscopically ;
and from any animal which looked in any way unhealthy cultures
were made.

The results were summarized as follows :—5 rats were caught
in the Park, and 495 in the Gardens: 283 of these were males
and 217 females.

3 rats had tubercle, 10 had tapeworm cysts in the liver, 49 had
Trypanosoma lewisi in their blood, 2 had empyema (not tubercular),
1 had a tumour of the lower jaw (the result of an old injury), and
1 had pleuritis and hydrothorax (not tubercular), ;

946 MR. R. I. POCOCK ON THE CRESTED RAT.

Bacteria were found in 71 rats: in 40 in the lungs, and in 31
in the spleen.

Saccharomycetes were found in the lungs of 16 rats.

Fleas were found on 4 rats, and lice on 3 rats.

The general condition of the rats was very good, and in none
was anything at all suspicious found.

Dr. R. W. Ssaurexpr, C.M.Z.S., sent for exhibition a photograph
he had taken of a living specimen of a male albino Woodchuck,
Arctomys monax, that had been sent to him from Virginia,

UWESsA;

Mr. R. E. Houpine exhibited and made remarks upon the
Horns of a Highland Ram, a Fallow Deer, and a Roebuck, which
were fused at the base, and also the skull of a coursing Greyhound
with abnornal dentition.

Mr. R. I. Pocock, F.R.S., F.Z.S., Superintendent of the Society’s
Gardens, exhibited the skin and skull of a specimen of the
rare Crested Rat, Lophiomys ibeanus Thos,, which had been sent
from Nakuru for the Zoological Gardens by Mr. R. B. Woosnam,
C.M.Z.S., but had unfortunately died on the voyage. After
alluding to the well-known peculiarities of the skull in this Rodent,
Mr. Pocock drew attention to the arrangement and coloration
of the hair (text-fig. 190), and expressed the opinion that the
alternating areas of black and white with which each individual
hair is ornamented, must make the animal conspicuous at dusk,
if confidence be placed in the analogy supplied by such animals
as Poreupines and Zorillas. The coat consists of two kinds of
hair,—a softer close-set under-fur dusky grey at the base, then
white, then dark at the tip, and much longer, coarser hair usually
dark at the base, then white, then black, then white at the tip.
These hairs are so arranged that the dark and white bands of the
nnder-fur coincide exactly with the alternating bands of the same
colour in the coarser hairs, while the long white tips of the latter
oroject clear of the rest of the coat. From this arrangement it
results that when the hair is raised the median white bands
combine to form a continuous white mass thrown into relief by
the dusky base and the distal black area, the whole being su-
mounted by the white tips of the long hairs shining with almost
silvery lustre.

The coat of the upper side, moreover, is divided into three
definite regions—a median dorsal and a lateral on each side—by a
band extending from the shoulder to the hip and consisting of
subspiniform greyish hairs of peculiar spongy texture, and thick
in the middle but narrowest at the base and apex. When the
animal is viewed from above with its coat erect, the white areas
of the region on each side below this dividing band form a

MR. R. I. POCOCK ON THE CRESTED RAT. 947

continuous white lateral stripe which anteriorly merges more or
less, according to the species, with the white area on the summit
of the head; and when seen from the side the same white stripe
is visible as well as the white area of the hairs of the dorsal crest.

Text-fig. 190. -

Dorsal and side views of Lophiomys ibeanus.

In the typical form of Lophiomys, namely LZ. imhausi, the tail
is longer and much whiter and the frontal band Fareeln longer
and more conspicuous than in ZL. ibeanus. ‘These two Fontes

948 THE HON, P, A. METHUEN ON

conjoined must, theoretically, combine to make the former species,
or race, more easily seen at night than the latter.

That the Crested Rat is nocturnal was proved by observations
made upon the specimen of L. wmhausi that was kept in the
Jardin d’Acclimatation, Paris. It was also recorded of this
animal by Milne-Edwards that when irritated it raised its dorsal
crest erect and defended itself by biting vigorously.

Mr. Pocock concluded by remarking that the specimen exhibited,
which before being skinned was perfectly fresh having been kept
in a refrigerator, had a most peculiar but indescribable smell.
From this it might be inferred that Lophiomys was perhaps a
protected self-advertiser. On the other hand, it was considered
possible that it might be a mimic of the Porcupine, since the
coloration of the two was in a general way very similar *.

PAPERS.

41, On an Amphipod from the Transyaal.
By the Hon, Paun A. Muruuen, F.Z.S.

[Received April 27, 1911: Read June 13, 1911.)

(Plates X LIX.—LI.*)

Introduction.

About twelve months ago, Mr. Hewitt, of the Albany Museum,
Grahamstown, who, at that time, was at the Transvaal Museum,
Pretoria, communicated in a letter to the Rev. Noel Roberts and
to me, his discovery of a blind Amphipod together with some
Copepods in a cave at Irene, which lies about 9 miles south of
Pretoria; at the time of writing he considered the species to
belong to the genus Hucrangonyx Stebbing. Owing to other
work he postponed the description of it, and, when I came out
some few months ago, he very kindly gave me the specimens
collected to deal with.

Not long ago I visited the same cave and secured two specimens :
none of the Amphipods taken from this cave were large.

In February, Mr. Austen Roberts and myself went to the
Makapan Caves, which lie in hilly country about 15 miles west of
Potgietersrust, in the Transvaal, for we heard that some of these
caves contained water, and we hoped to obtain a more plentiful
supply of these cave Crustacea.

The first cave we visited—the more famous historically—con-
tained nothing we were in search of, so we directed our steps to

* Tn the discussion that followed this exhibition, Dr. R. E. Drake-Brockman, who
was acquainted with Lophiomys in Somaliland, remarked that the natives of that
country regarded these rats as young Porcupines.

«-+ Foy explanation of the Plates see p. 957.

“ayo URUTMAL '}S3AQ

IsSbaHeod

XANODNVAEONH

Ie ies) dlchiltle Jeu Ti,

.
P
j
4
A

West,Newman lith.

BUCRANGONYX ROBERTSI.

PGS) eS) JIL LAL

oe ras
eas

Zi,

iy

West,Newman lith.

BHUCRANGONYX ROBERITSI.

AN AMPHIPOD FROM THE TRANSVAAL, 949

two others on the opposite side of the valley. In both we found
a plentiful supply of water.

As to these two caves, one was considerably deeper than the
other. In the first one we entered, we reached the level floor
fairly soon ; here we found shallow sheets of water supporting a
large number of Gammarids, Copepods, and Ostracods ; the bottom
was muddy and the floor of the cave generally covered by the
droppings of bats. No light entered as far in as this. The
bottom of the second cave we reached after a rather long, steep,
and winding descent: the stalactites and stalagmites showed it
to be a limestone cave. The water here was as clear as crystal,
rich in Gammarids and poor in Copepods; there were no
Ostracods, but a few aquatic worms were taken; some terrestrial
Isopods and some spiders were also found. Though the water
appeared quite still, a fresh supply was without doubt being added
continually. The bottom was rocky, covered by a thin layer of
fine mud, and the floor of the cave, as in the other, was strewn
with bat droppings.

Later in the day a few Gammarids, similar in every respect
to the others taken, were caught under stones at a spring in
the vicinity, but none were ever taken in the spruit which runs
through the valley, where the numerous crabs would make short
work of such fry.

All the Gammarids so far found in the Transvaal represent a
single species, belonging most probably to the genus Hucrangonya
Stebbing, and closely related to Hucrangonyx vejdovskyi Stebbing,
vide (3. p.389 and 4). However, this little cave form has not been
placed in this genus without notice being taken of certain resem-
blances and affinities to the genus Viphargus Schiddte(3. p. 405),
more so to Weoniphargus Stebbing (3. p. 404 and 2, p.73), and
the genera Crangonyx Bate and Paracrangonyx Stebbing (8.
p. 969 and 1. p. 218). In many ways this species appears to be a
generalized Niphargus-Crangonyx type, aS 1s seen mainly in the
nature of the telson and third uropods ; the breadth of the second
joint of pereiopods 3 to 5, and the total length of these appendages
as compared to pereiopods 1 and 2; the shape and size of the
gnathopods ; the number of sete and spines on the inner and
outer plates of the maxillula; the structure of the lips; the size
of the accessory flagellum and total length of the antennule.

It is in the sense of the comparative generalization of its
structure that this creature may be called primitive; primitive as
opposed to the more recent genera Paracrangonyx, Apocrangonyx,
and Crangonyx, but, on the other hand, more recent than the
genus Gammarus, and probably NViphargus.. This conclusion has
been arrived at more by the study of the telson than of any other
part (vide 1. p. 219).

The large size of the outer ramus of the third uropod appears
to be almost peculiar, but, judging from Chilton’s (1. p. 218)
remarks on the variability of this element in Crangonyz, this
character can be given apparently too much attention.

Proc. Zoot, Soc.—1911, No. LXV. 65

950 THE HON. P. A. METHUEN ON

It may be as well to record the outstanding characters of this
new species in order, and mention points of similarity and dif-
ference between it and the various genera and species mentioned
above, as they appear significant in leading us to some conclusion
as to the proper systematic position of the only known fresh-water
Amphipod from South Africa (unless some littoral marine forms,
taken in a stream in the Cape Peninsula and in vleis on the Cape
Flats within two or three miles of the sea, be considered as true
freshwater creatures).

The antennules are longer than the antenne, the accessory
flagellum small and 2-jointed, subequal to the first joint of the
normal flagellum. In Hucrangonya vejdovskyi, the second joint
of the accessory flagellum is half the length of the first. In
Neoniphargus, the accessory flagellum is 2-jointed, longer or
shorter than first flagellular joint.

Upper lip rounded as in Viphargus, Crangonyzx, etc.

Lower lip, inner lobes fairly small; in Paracrangonyx they are
small; in Wiphargus well developed; in Crangonyx flagellula
Benedict, very small. Mandibular processes moderate; in Hucran-
gonyx vejdouvskyi unusually prolonged.

Mandibles normal, dissimilar. Second joint of palp, subequal
to third, is rather broad owing to convexity of inner margin; in
Eucrangonyx vejdovskyi second joint broad, considerably longer
than third; in Paracrangonyx third joint subequal to second,
which is not expanded (in figure) ; in Viphargus third joint longer
than second.

Maxillula : inner plate with two plumose setz, outer with seven
spines serrated on the inner side; two or three sete to inner
plate in Niphargus and Neoniphargus, the former with seven
spines to the inner plate, the latter with six ; in Paracrangonyx
the same as for Hucrangonyx robertsi; inner plate of Crangonyx
with six sete, of Hucrangonyx vejdovskyi with four setee.

Maxilla.as in Hucrangonyx.

Maxillipeds as in Hucrangonyx (most probably), not unlike
those in Paracrangonye.

The two pairs of gnathopods as in Hucrangonyx vejdovskyt, but
palm equal to hind margin; in #. vejdovskyi it is much shorter.

Pereiopods 1 and 2 slender and slightly shorter than succeed-
ing pereiopods; in H. vejdovskyi they are “said to be slightly
longer.” Pereiopod 5 the longest. The second joints of all the
pereiopods much as in Z. vejdovskyi.

Pleopods normal; in Paracrangonyx slight and one-branched.

In uropod 1, peduncle a little longer than the equal rami; in
E. vejdovskyi as long.

In uropod 2, rami slightly unequal, peduncle as long as shortest
ramus; in #. vejdovskyi shorter than rami.

Uropod 3 has peduncle half the length of long outer ramus,
inner ramus minute with single spine; in H. vejdovskyi peduncle
half the length of outer ramus, inner ramus flattened, rudi-
mentary, shorter than peduncle; in Crangonyx and Paracran-

AN AMPHIPOD FROM THE TRANSVAAL. 951

gonyx uropod 3 not elongate, so also Neoniphargus which has
inner ramus minute; Miphargus has outer ramus long, two-
jointed, and the inner small.

Telson with emargination and almost square, as in ZL’. vejdovskyi ;
in Crangonyx entire or partly cleft; in Paracrangonyx entire ;
in Veoniphargus partly cleft ; in Viphargus deeply cleft.

This evidence seems sufficient to warrant the inclusion of
this new species in the genus Hucrangonyx near the species
EL. vejdovskyi.

I have taken the opportunity here offered of naming the species
after the Rev. Noel Roberts, in recognition of his enthusiasm for
this branch of zoology.

Detailed description of the new species.

Family GAMMARIDA® Leach.
Genus Eucranconyx Stebbing.

EUCRANGONYX ROBERTSI, sp.n. (Plate XLIX.)

Female: length of largest specimen taken 11 mm.*; colour
dirty yellow or pink in the shallower cave near Potgietersrust,
semi-transparent white in the deeper cave. No trace of eyes
was discovered.

Body rather compressed, smooth, no carina, rostrum barely
perceptible, a few minute spines on dorsal part of head and third
segments ; the last three or five segments with spines on posterior
margin.

First four coxal plates of pereion deeper than the rest, the first
smallest, the third and fourth deepest ; the fourth plate broader
than the others; side-plates of the fifth, sixth, and seventh
pereion segments small, those of fifth and sixth deeply emarginate
behind. These coxal plates bear marginally small spines.

Antennules (Pl. L. fig. 1) long, longer than the antenne;
flagellum much longer than the peduncle; first joint of the
peduncle a little shorter than second and third joints together ;
second joint rather longer than third ; accessory flagellum (Pl. L.
fig. 1 q@) two-jointed—both joints armed with sete,—minute,
hardly as long as first joint of flagellum; all the joints of the
antennule provided at their distal extremity with moderate sete,
of which one or two on each segment are conspicuously longer
than the rest; the second joint of the peduncle has also two
groups of small setee and one other seta besides about its middle
length ; the arrangement of the setz on each joint of the flagellum
appears to be constant in disposition after the eighth joint, and
to be about seven in number.

Antenne (Pl. L. fig. 2) about four-sevenths the length of the

* The length includes that part from the anterior margin of the head to the
posterior margin of the last pleon segment.
65*

952 THE HON. P. A. METHUEN ON

antennules; peduncle longer than flagellum, the ultimate and
penultimate joints well armed with numerous sete which are
arranged, generally speaking, in succeeding and nearly parallel
semicrescents about all parts of the joints except the proximal
extremities; on the other joints of the peduncle the sete re-
present a much diminished quantity; the distal parts of the
flagellum joints carry about eight sete each, disposed in an un-
varying sequence.

Lips: wpper lip (Pl. L. fig. 4) rounded, with some minute setz
about the extremity, at the centre of which they take an inward
course for a short way on each side. Lower lip (Pl. L. fig. 3):
inner lobes with minute sete on distal convex margins; outer
lobes with longer setze; mandibular process not elongate, bearing
small sete.

Mandibles (P\. L. figs. 5 & 5a): palp rather long when com-
pared with the size of the mandible itself; first joint shortest,
devoid of sete; second joint rather broad and curved, the side
opposite the biting process markedly convex and armed with
two rows of sete (about fifteen in number); the ultimate joint is
about the same length as, or a little longer than, the penultimate
and is provided with terminal sete on both lateral margins. Palp
on each side similar except that basal joints are of slhghtly
unequal lengths. Mandibles themselves of unequal size and of
slightly different structure (vide figs. 5, 5 @).

Mawillula (Pl. L. fig. 6) resembles that of Paracrangonyx com-
pactus Chilton very closely. The first jomt of the palp is short,
the terminal joint bears about nine bristles; the endite of the
third joint of the maxillula is armed with seven stout serrated
bristles, which resemble those found in a similar position in
P. compactus but differ in being more extensively serrated, and,
further, at the base of the innermost of these bristles is found a
group of fine hairs; this endite is about double as broad as the
palp; the endite of the first maxillula joint (accepting Hansen's
interpretation of the endites) appears to be exactly similar to
that of P. compactus, bearing two fairly long plumose sete and a
number of hairs on its inner margin.

Mazilla (Pl. L. fig. 7): the endites are simple, regular, and
well-developed structures, both armed distally with a great number
of setee which are disposed in two rows, the more distal of these
rows being considerably longer than the rest, at any rate in the
case of the endite of the third jomt; the inner side of the endite
of the second joint bears a number of fine hairs, and between
these and the distal rows of setze les one long plumose and another
smaller, slightly plumose seta.

Mazilliped (P1. L. fig. 8): the shape of the joints like those of
Paracrangonyx compactus, the whole limb being, however, some-
what slenderer ; the seventh joint alone is shorter than that of
P. compactus; a great number of closely-set sete and bristles
clothe the inner surface of the two endites and of the fifth, sixth,
and seventh joints. ‘The basipoditic endite or inner plate carries

AN AMPHIPOD FROM THE TRANSVAAL. 953

three plumose sete on its inner margin, and distally three short
stout bristles and about eleven plumose setee. The endite of the
ischiopodite or outer plate carries a number of fairly short stout
bristles distally, and along two-thirds of its inner margin six of
the distal bristles are serrated.

Gnathopods: about the same size and length; the fifth and
sixth joints of gnathopod I. somewhat broader than those of
gnathopod II. In the male, gnathopod I. is a little longer than
gnathopod IT.

Gnathopod I, (Pl. L. fig. 9) well developed, subchelate; the
coxa of the first joint in shape making almost a parallelogram,
sloping forward somewhat, anterior portion deeper than posterior,
diagonal from posterior ventral corner to anterior dorsal corner
equalling greatest length of coxa; anterior margin bearing five
small spines and ventral margin five also. The second joint or
basos is narrow at the base, has anterior side straight, and bears
six feebly serrated sete, the most distal the longest; posterior
margin convex, most markedly about the middle of the joint
which is here seen to be two-fifths in breadth to length; the
posterior margin carries a number of long sete, and following
this arrangement, four setze proximally, equidistant apart, followed
by a group of two, then a group of three, a group of two, another
group of three setz all serrated, and distally a group of four
smaller sete, not serrated. The third joint or ischium is short,
broader than long, and bears on its posterior distal extremity five
sete not serrated. The fourth joint or meros has the posterior
distal portion rounded and well provided with numerous serrated
setee, proximal to which lie a group of four serrated sete; on the
anterior margin four small sete can be noticed. The fifth joint
or carpus, which is more or less triangular, has a dense cluster of
serrated setze on its short posterior portion, which are arranged
into groups, one consisting of a row of marginal sete, another of
apparently two rows placed submarginally ; the anterior margin
of the joint carries four sete. The sixth joint or propodos is
subovate, equal in length to the longest measurement of the basos,
as wide as the greatest vertical depth of the coxa; anterior margin
convex, with four groups of four fairly short sete arranged at
varying intervals from a fifth group of six longer sete situated
near the base of the dactylos; in addition, a small solitary seta
will be noticed external to the third of these groups; the posterior
margin, starting from the proximal end, is straight for a short
distance, to a part where four small well-marked prominences are
encountered ; on each of these are placed two finger-shaped pro-
cesses of unequal length and, excepting on the last prominence,
two sete ; behind the fourth prominence lies a row of sete: the
palm, to which the dactylos fits closely, follows immediately ; it is
provided with a series of very short spines arranged in couples,
each couple equidistant apart; a number of short sete, nine in
number, are also present opposite the border of the most posterior
muscle, disposed more or less in a straight line, and varying

954 THE HON. P. A. METHUEN ON

slightly in individual length ; between the eighth seta of this
series and the middle portion of the palp are placed three fairly
long sete, of which the central is the longest. The dactylos,
which is slightly curved, is provided with two sete on its anterior
and four very short sete on its posterior margin; this inner
margin bears distally a small tooth distinct from the long
curved terminal unguis; between these two lie two small sete.
None of the sete on the ultimate and penultimate joints are
serrated.

Gnathopod II. (Pl. LI. fig. 10) is very like gnathopod I. Only
the chief differences will be mentioned. The sete on the second
joint are somewhat different in disposition and in length. The
third joint is somewhat longer and narrower and possesses more
setee. The fourth joint has fewer sete and only one fairly dense
group of about ten sete. The fifth joint is three-quarters as
broad as long (the same measurements in gnathopod I. show the
breadth to be greater than the length); the anterior and posterior
margins are comparatively long, the former the longest; the sete
on the posterior margin are longer and more numerous and are
arranged in seven or eight groups. The sixth joint has, on the
anterior part, an additional group of fairly long sete, which vary
in number in each group: the whole joint is longer and narrower:
the palm does not differ much, but the proximal half of the pos-
terior margin bears eight groups of long feebly serrated setee,
between which and the palm lie five finger-shaped processes, one
conspicuously longer than the rest with another little one at its
base. The dactylos and terminal unguis spine are both shorter ;
between the latter and the small terminal tooth there are no
sete.

Pereiopod J. (Pl. LI. fig. 11): the coxa of the first joint is
almost rectangular; the anterior and ventral margins are,
however, somewhat concave, the middle part of the posterior
margin slightly convex; the anterior margin has four short sete,
the ventral three. The second joint is long, narrow at the
proximal base, whence it widens rapidly but nowhere greatly,
being never more than a quarter as broad as long; nine spines of
moderate length and seven spines of greater length arm its
anterior and posterior margins respectively ; further, a group of
four sete are placed at the distal extremity of the posterior
margin. The third joint is a little longer than that of the
gnathopods but is considerably narrower; its posterior margin
carries a group of four setz distally. The fourth joint is long,
longer and broader than the fifth, the two joints together about
the same length as the second joint: the anterior margin carries
three setee unequal in length—at the base of the longest a minute
seta can be seen—and distally a group of three sete, one of which
is long; the posterior margin bears three groups of three sete
each, then one short seta, then distally a group of three sete.
The fifth joint, which is equal in length to, but broader than, the
sixth joint, carries on its anterior margin two minute sete, and

AN AMPHIPOD FROM THE TRANSVAAL. 955

distally a group of three small sete; the posterior margin is pro-
vided with four groups of sete, the proximal group consisting of
three sete, the next of two, then another of two, the most distal
group of four medium and one large seta. The sixth joint, which
is slender, bears on its posterior margin six groups of three sete
to each group, and on its anterior margin three minute sete, and
distally a group of four setze.

Pereiopod II. is practically similar in all respects to pereiopod L.,
to which it is subequal; on the second and third joints there are
two or three small additional sete marginally.

Of the remaining pereiopods the fifth is the longest, the third
the shortest, these differences of length being appr eoable but not
great; they are, however, considerably longer than the first and
second. In the third pereiopod, although the second and third
joints are somewhat shorter than those of the first pereiopod, the
great difference in length is made up by the much greater length
of the fourth, fifth, sixth, and seventh joints of the third pereiopod;
at the same time these four joints, though slender, are of the same
breadth as those of the first pereiopod.

Pereiopod II], (Pi. LI. fig. 12): the coxa of the first joint is
somewhat concave anteriorly, emarginate posteriorly ; anteriorly
and ventrally it bears some small spines. The second joint is
rather narrow at the base, broadens very rapidly, and then
narrows gradually towards its distal extremity ; it is a little more
than twice as long as broad; the anterior margin is slightly
convex and is beset witha number of moderate spines—deseribing
from the proximal end, three single ones equidistant apart, six in
pairs subequidistant apart, and terminally a group of three spines:
the posterior margin has fourteen shorter spines equidistant apart
and terminally two closer together; behind these latter a large
spine. The third joint is slightly rounded and carries a group of
four fairly small sete. The fourth joint is broader at the base
than that of the first pereiopod; the anterior margin carries nine
small sete, arranged in pairs all but the most proximal, and
distally a group of four sete, one conspicuously larger than the
others; the posterior margin carries two moderate spines and
three small ones and distally two fairly large and two small spines.
The fifth joint is armed with five groups of spines, the most
proximal group of two, the next three of three, and the following
of two spines to each group, and distally with a sixth group made
up of two larger and two smaller spines; opposite the last group
is a group of one larger and three smaller spines, some distance
behind which is a single spine, and behind this spine again two
groups of three spines, and disposed proximally to these a couple
of small spines close together. The sixth joint is conspicuous on
account of the Jarger size of some of the spines on its anterior
margin ; on this margin four groups of three spines to each are
fixed, and distally a group of eight spines varying in length:
the opposite margin six groups of spines are seen, each ee
possessing two or three spines. The seventh joint, including the

956 THE HON. P. A. METHUEN ON

claw, is one-third as long as that of pereiopod I., and carries on
the posterior margin two small spines.

In pereiopod IV. the details of the arrangement of the sete
differ very slightly.

In pereiopod V. the third and fourth joints are equal in length
to, but broader than, those of the third pereiopod; the seta
arrangement is not exactly similar, but this slight difference is
negligible. In Irene examples the penultimate joint of this
appendage carries much longer spines on one side than is the case
in specimens from the Makapan caves.

The pleopods are normal; the basal portion bears two branches
and two little coupling-hooks. The longest pleopod is the
second.

Pleopod I. (Pl. LI. fig. 13): basal portion oblong, slender, and
devoid of sete, except the two coupling-hooks ; these little
structures bear a few tubercles and one slightly hooked at the
end. The inner ramus is somewhat longer than the outer; each
ramus consists of thirteen joints which bear long plumose sete.

Pleopod II.: the basal part bears distally on its anterior margin
two fairly long spines, which are absent in the other pleopods and
appear to be absent altogether in the male.

Pleopod III.: the number of joints to the rami are two or
three less than those of the first pleopod.

The wropods: the first uropod the longest; the third uropod
not quite so long as the first; the second uropod about two-thirds.
as long as the first. The peduncle of the first uropod long, longer
than either ramus, much longer than peduncle of second; peduncle
of second much longer than peduncle of third. Outer ramus of
third uropod long, much longer than those of first ; inner ramus
minute. Rami of first uropod longer than those of second.

Uropod I. (Pl. Li. fig. 14): peduncle, broadest at the base,
bears five spines on posterior margin, and two others close
together distally. Outer ramus a trifle shorter than inner, which
bears five spines on posterior margin and two larger and four
smaller ones at the extremity. The outer ramus with four rows
of spines, three in each row, and another row basally of two
spines; distally two larger and two smaller spines.

Uropod IT, (Pl. Li. fig. 15): peduncle fairly stout as compared
with that of uropod I., narrower basally, but broader distally than
same; on posterior margin two spines; on median ridge (vide fig.)
five spines, the most distal being the longest. Inner ramus
longer than outer, which carries on posterior margin four spines,
and distally two longer and five shorter spines; anterior margin
has two small spines; the outer ramus bears posteriorly four
spines, two longer and three shorter distally, and eight spines
arranged in couples anteriorly.

Uropod ITI, (Pl. LI. fig. 16): peduncle about the same breadth
as that of uropod IJ.; anterior margin bears distally two small
spines; opposite these is seen the inner ramus bearing one small
spine, not counting spine it is one-fourth the length of the peduncle.

AN AMPHIPOD FROM THE TRANSVAAL. 957

Anterior margin of outer ramus carries six groups of spines, three
to each group except the two most distal, which are made up of
two spines each; distally is a group of fairly long setz: the
posterior margin shows five groups of spines, the first two groups
of three spines, and the other three of four spines each. At
junction of peduncle and outer ramus three spines can be dis-
tinguished. In the male the outer ramus appears to be hardly as
large as in the female.

Telson (Pl. LI. fig. 17) almost square; sides slightly convex :
two small plumose sete on each side bear two larger and two
smaller spines placed in a posterior position. The structure
excavate behind; in the male this excavation not quite so deep
as in the female. In specimens from Irene one of the terminal
spines on each side is much longer than in other specimens.

Literature referred to.

1. Cuinron, C.—‘ The Subterranean Crustacea of New Zealand.”
Trans. Linn. Soc., 2nd ser. Zool. vol. vi. pt. 2, 1894.

2. Smirn, G. W.—“‘The Fresh-water Crustacea of Tasmania.”
Trans. Linn. Soc., 2nd ser. Zool. vol. xi. pt. 4, 1909.

3. Srepsrne, T. R. R.—Amphipoda Gammaridea in Das Tierreich,
21 Lief. Berlin, 1906.

4. Vuspovsky.—Crangonyx subterraneus (non Bate 1859) in SB.
Bohm. Ges., nr. 10, p. 12, t. 1-3.

EXPLANATION OF PLATES XLIX.-LI.

The appendages have been drawn with the aid of a camera lucida.

All the figures are of Hucrangonyx robertsi, 2.

PratE XLIX.
Side view of female, X 11.

Prate L.

Fig. 1. Antennule, x 29.
la. Accessory flagellum, X ca. 160.
2. Antenna, X 29.
3. Lower lip, X ca. 45.
4, Upper lip, X ca. 45.
5 & 5a. Mandibles, X 29.
. Maxillula, X 75.
. Maxilla, X 75.
. Maxilliped, X 40.
. Gnathopod I, X 29.

co O-1

Prats LI,

Fig. 10. Gnathopod IT, X 29.
11. Pereiopod [, x 29.
12. Pereiopod III, X 29.
13. Pleopod I, x 29.
14. Uropod I, X 29.
15. Uvropod II, x 29.
16. Uropod III, x 29.
17. Telson, X 29.

Text-fig. 191.

958 MR. R. LYDEKKER ON

42. An African Rhinoceros, Klipspringer, and Gazelle *.
By R. LyprexKer f.

[Received May 9, 1911: Read June 13, 1911.]
(Text-figures 191-193.)

I. THe Somatt RHINOCEROS.

The presentation to the British Museum by Mr. Drake-
Brockman of two skulls of the Somali Rhinoceros, affords an
opportunity of considering whether that animal is entitled to
rank as a distinct local race of the so-called black species.
Sportsmen, I am told, almost invariably regard it in that light;

* The complete account of the two new forms described in this paper appears
here, but as the names and preliminary diagnoses were published in the ‘Abstract,’
the species are distinguished by the names being underlined.—Ep1Tor.

+ By permission of the Trustees of the British Museum.

Upper aspect of skulls of East African (A) and Somali (B) Rhinoceroses,

THREE AFRICAN MAMMALS. 959

and Mr. Ward informs me that such heads as he has mounted
indicate a relatively small animal, with horns inferior in size to
those of the Eastern and the Southern Rhinoceroses, and a skin
with a somewhat different epidermal structure.

According to modern views in regard to nomenclature, the
Somali Rhinoceros already has a scientific name, since it is re-
ferred to by Count Joseph Potocki on page 82 of his work entitled
‘Sport in Somaliland, London, 1900, as Rhinoceros bicornis
somaliensis; and although no description was published at the
time, the accompanying plates apparently render the name valid.

Text-fig. 192.

Front (A) and side (B) views of skull of Nigerian Klipspringer.

Of the two skulls presented by Mr. Drake-Brockman, one is
that of a subadult animal, with the whole of the permanent
dentition in use, and almost perfect, although the tip of the pre-
maxille is broken off. The other, which is considerably more
imperfect, is that of a younger animal, with the upper premolars
only just coming into wear.

Compared with a skull of Rhinoceros bicornis from East Africa
(B.M. No. 7.2.26.1)* (text-fig.'191), of somewhat greater age than
either of the Somali specimens, the latter are seen at once to
differ by the narrower form of the whole upper surface, both at
the interparietal constriction and at the orbital expansion. The
boss for the front horn is also much less expanded in the Somali
skulls, and there is less convexity in the region immediately behind

* T am fully aware that this is not the type locality of the species.

960 MR. R. LYDEKKER ON

this. Moreover, the palate is more decidedly vaulted in the
Somali than in the East African skull. So faras I can ascertain,
these differences appear to be constant in all the skulls available
for comparison.

The differences in the proportions of the Somali and East
African skulls will be apparent from the following table :—-

E. African, Somali.

iKensbh ofjupper aspectiwmsaeeceenceteee ee eens 227 ins. . 23 ims.
reali hsabhon outs aieee anne see ee anne 11 94
TPRVEMRAIUG VAIN Gbegysoc 200 cccaaansormonssassean eas. 14 203
A ROTMEM HO WANCNE D5 45 sco opgss0gedgo0nsoooobosceoneseS 142 Te
Length of upper tooth-row (excluding p.1)... 102 10¥
AW elit has Sot ain’ Ss cas, Ns hone ate et eee Pac a 24

As these dimensions amply demonstrate the racial distinctness
of the two forms, the Somali animal may stand as &. bicornis
somaliensis Potocki; the specimen here described occupying the
position of type. It may be added that if the East African and
Somali skulls were of the same age, the difference in the lengths
of the upper tooth-row would be greater.

Ii, Tue Nicrrian Kirpsprincer (Oreotragus saltator porteust).
Lydekker, Abstract P. Z.S. 1911, p. 38 (June 20).

Early in May, as I have already stated in the Field news-
paper for that month, Mr. Rowland Ward directed my attention
to the skull and borns of a male Klipspringer from the Duchi ’n
Wai range of the Yola province of Northern Nigeria, lying to
the south-west of Lake Chad. The skull was taken from an
animal shot there by Dr. E. J. Porteus, by whom it was kindly
piaced at my disposal. Klpspringers, it appears, are quite
familiar to the natives working in the Yola tin-mines, by whom
they are known as gaddi-dueki, a term equivalent to Hill Duiker,

The skull (text-fig. 192) differs from that of an East African
Klipspringer by its much greater width ; its diameter across the
orbits being 34 inches, whereas that of the Hast African specimen
is 212 inches, It is further distinguished by the marked bending
down of the margin of the lachrymal bone, which consequently
has a distinct lateral surface in place of a sharp edge, and also
shows only a comparatively small part of its total area from the
frontal aspect.

Klipspringers, so far as [ am aware, appear to be unknown on
the West Coast north of Angola*; while to the north, Abyssinia
is the nearest point to Yola where they are found. As the
country between the last two districts is to a great extent low
desert, the Yola Klipspringer must be quite isolated.

On this ground, and also on account of the peculiarities in the
form of the skull, it is clearly entitled to racial distinction, and
I propose to name it Oreotragus saltator porteusi, in honour of
Dr. Porteus.

* For the information that Klipspringers inhabit Angola, I am indebted to
Mr. E. A. Hamilton.

THREE AFRICAN MAMMALS. 961

The type of the new race of Klipspringer will be the aforesaid
skull, which Dr. Porteus has kindly presented to the British
Museum.

I know of no Antelope with a distribution identical with that
of the Klipspringer, as now extended.

About a fortnight after describing this skull I received a letter,
dated Naragata, Northern Nigeria, from Mr. M. P. Hyatt,
informing me that he had recently killed three Klipspringers—
an adult buck and two does—in that part of the country.

IIT. An ALGERIAN GAZELLE (Gazella hayi).
Lydekker, Abstract P. Z.S. 1911, p. 38 (June 20),

At the close of 1909, Mr. M. V. Hay presented to the British
Museum the skin and skull of an adult male Gazelle, shot by
himself in Algeria between Constantine and Biskra, and sup-
posed to be a Doreas (Gazella dorcas). In due course the specimen
was set up, and placed on exhibition as a representative of that
species. Recently, however, as already mentioned in the Field
newspaper *, it was pointed out to me by the donor that the
specimen differed considerably from the Dorcas, and I was informed
at the same time that its distinctness is reeognised by the Arabs,
who call it rhozal-rim ; rhozal being the name of the dorcas, and
rim that of Loder’s Gazelle (G. leptoceros). On comparison of the
specimen with an undoubted Dorcas from the Biskra district, the
difference between the two became apparent (text-fig. 193).

Mr. Hay’s specimen is about the same size as a Dorcas, but
appears to have rather larger ears,—I say appears to have, because
one 1s never certain whether there may not have been stretching
or shrinking in the mounting. Its most distinctive characteristic
is, however, to be found in the form of the horns, which com-
pletely lack the double, sublyrate curvature of those of the Dorcas,
and incline almost directly upwards and slightly inwards, with a
slight inward and forward turn at the tips. There are also much
fewer rings on the horns, the present specimen having only 12,
whereas a Dorcas may have 24 or 25; possibly a more aged
example of the new form might develop two or three more rings,
but even then the difference would be very considerable.

The face-markings differ considerably from those of a Dorcas,
and are more like those of an Edmi (4. euvieri), the middle stripe
being much darker, with a conspicuous nose-spot, and the light
eye-stripes much less apparent, while the forehead lacks the
chestnut tint of the Dorcas. The body does not show the faint
light stripe above the flank-band which occurs in the Dorcas ; the
knee-tufts are larger and blacker; and the tail is brown instead
of black, with a shorter fawn area at the base.

As the skull of Mr. Hay’s specimen is mounted in the skin, I
cannot give cranial characters; but the foregoing features are
amply sufficient to distinguish the new Gazelle from the Dorcas,

* June 3rd, 1911.

962 ON THREE AFRICAN MAMMALS.

which is found much more abundantly in the district: frequented
by the former. ‘This being so, there is no doubt as to the dis-
tinctness of the new species, which I have named Gazella hayi,

from all other Algerian Gazelles, unless it be G’. kavella and the
so-called Antilope corrina, neither of which can now be identified.
The type of the new species will, of course, be the Museum
specimen.

both from the Constantine—Biskra district,

t)

Heads of Gazella hayi (A) and G, dorcas (B)

P49 Ili, Jl, ILI.

Witherby & Co., Imp.

CAPRA PYRENAICA HISPANICA.

(Summer Pelage)

AS Iii, 2, Li.

Witherby & Co, Imp.

CAPRA PYRENAICA VICTORIA 4é, &.

(Summer Pelage)

hye al
:

ae

IP ZoS> t@iil, Pi, ILM

Witherby & Co., Imp.

CAPRA PYRENAICA VICTORI€.
(Winter Pelage)

ON THE SPANISH IBEX. 963

43. The Subspecies of the Spanish Ibex.
By Prof. Ancen Caprera, O.M.Z.S.

[Received May 13, 1911: Read June 13, 1911. ]}
(Plates LIT.-LIV.* and Text-figures 194-199.)

The first scientific description of a Spanish Ibex was that
published in 1833 by F. Cuvier, in the great iconographic work
‘ Histoire Naturelle des Mainmifeéres,’ pl. 396. It was based on
a young male from the Pyrenees, of which the author gave
a beautiful figure in winter pelage, but was not accompanied by
any technical name. In the index of the work, published in 1842,
the animal is erroneously called Capra ibex, the fact being
apparently ignored that four years before Schinz had described
the same forn m of Goat under the name Capra p yrenaice T ie

In volume xxvi. (1848) of the ‘Comptes Rendus’ of the Paris
Academy of Science, Schimper briefly mentioned the Ibex of the
Andalusian sierras, naming it C. hispanica and considering it as
quite a different species, a view followed by all the authors “of the
time, and sustained still in our own days by F orsyth Major =
and Graells$. Modern zoologists, however, seem to agree in
admitting only one species of Spanish Ibex, though admitting that
there are some differences between the specimens coming from the
Pyrenees and those from other parts of the Peninsula. Sclater ||
considers the latter as a “ slightly altered phase” of the foruione
and ‘Trouessart, in his ‘ Conspectus Mammalium Europe,’
expresses the same idea in a more modern fashion, describing
two different subspecies: Capra pyrenaica, from the ‘“‘ chaine des
Pyrénées,” and C. pyrenaica hispanica, from the “chaines de
montagnes de |’Espagne Centrale et Méridionale.”

I do’ not intend to discuss now the meaning of the terms species
and subspecies. While awaiting a satisfactory and univer. sally
accepted definition of these wor ds, I agree with other authors in
considering all the Spanish Wild Goats as belonging to a single
species ; but as to the number of subspecies, I think there are,
not two, but three at least, the Ibex of Central Spain being quite
different from both the Pyrenean and the Andalusian forms.
This view has been anticipated by Ménégaux in Perrier’s ‘ Vie
des Aniimaux,’ as he says that ‘la forme qui habite les sierras du
centre de la Péninsule fait le passage entre les deux formes
susnommeées ” (pyrenaica and hispanica). This central subspecies
remains, however, unnamed and undescribed as a different form,
and to name and describe it are the chief purposes of the present.
paper.

* For explanation of the Plates see p. 977.
+ Neue Denkschr. Allg. Schweiz. Ges. Nat. ii. 1838, p. 9.
t Atti Soc. Tose. Sc. Nat. iv. 1879, p. 2.

Memorias Acad. Cienc. de Madrid, Xvil, 1897, p. 356.
|| P. Z.S. 1886, p. 315,

964 PROF. A. CABRERA ON

Before doing so, must remark on the geographical distribution
of the species and on the differences between the various races.

In that interesting book ‘ Unexplored Spain,’ MM. Chapman
and Buck have quite recently told the history of the Wild Goats
of Spain and the manner in which they are protected against
imminent extinction, giving some details about their present
distribution. The authors are not exact, however, in saying that
the isolated colonies now formed by the Ibex have been “ separated
from each other during ages.” There are, in fact, strong reasons
for believing that in the past Ibexes inhabited every suitable
point of almost every mountain ridge in Spain. Names recalling its
existence, suchas Las Cabras, Cabrales, Cabrera, Cabreira, Cebreros*,
&e., are quite common in all the mountainous districts. In the
seventeenth century the species was found in all the Sierra Morena
and Sierra de Cazorla. The Sierra de Segura, Sierra de Francia
in the Salamanca Province, and the Toledo Mountains, where it
does not exist to-day,’formed parts of its range sixty years ago,
and it has been found in the Sierra de Bejar, between the Sierras
of Francia and Gredos, so recently as 1897 7; and in 186], the
date of Seoane’s ‘ Fauna mastolégica de Galicia,’ a few individuals
remained in the mountains of that region. That Ibexes inhabit,
or at least inhabited in 18904, the mountains of Gerez, in the
northern border of Portugal, is a well-known fact.

In connection with the existence of Ibexes in the western
extreme of the Cantabrian chain, it must be remembered that
fossil remains of Capra pyrenaica have been found in Santander,
and from this we may surmise that the species reached the north-
western corner of the Peninsula from the Pyrenees, through that
northern ridge. It spread from there southwards into the
central sierras, either through Portugal by the Serra da Estrella,
included in its range by Trouessart, or directly through the
Burgos mountainsand the Guadarrama. Father Saturio Gonzalez,
a noteworthy collector of mammals, tells me. that he has found in
the old monastery of Silos a pair of Ibex horns which have been
preserved there for centuries, and about sixty miles east from
Santo Domingo de Silos there are a Sierra de Cabrejas and
a village named Cabrejas del Pinar. ‘These facts seem to suggest
that the Ibex was once common in the mountains connecting the
northern chain with the Sierra de Gredos, but, since there is not
any evidence of its existence in the Guadarrama, too much
confidence cannot be put in this hypothesis.

As to the Andalusian sierras, it is evident that the Ibex found

* Oebra was frequently used instead of cabra as a name for the Ibex during the
twelfth and thirteenth centuries. It was from this that the old Spanish writer
Father Sarmiento supposed that Zebras formerly existed in Spain. The modern
vernacular naine of the animal is cabra montés (Mountain Goat), or simply montés,
but there are some other local names. In the mountains about the lower Ebro it is
called sawvatge (the wild one); the people of Galicia call it.craba brava or craba fera
(Wild Goat), whereas in the Pyrenees the name bucardo, related to the English buck
and the French bouwe and bowqguetin, is commonly used.

+ Rivas Mateos, Actas Soc. Esp. Hist. Nat. 1897, p. 208.

+ P. de Oliveira and L. Vieira, Annaes Scienc. Naturaes, 111. 1896, p. 91.

Text-fig. 194.

THE SPANISH IBEX. 965

its way to them by the Toledo mountains and the Sierra Morena,
turning afterwards along the mountainous ridges of Hastern
Spain northwards to the lower Ebro basin. Notwithstanding the
proximity of this region to the Pyrenees, I cannot believe that
the Ibex inhabiting it immigrated directly from the Pyrenean
chain, as it is very different from the form found here, and quite
indistinguishable from the southern race.

&

Baceanic !s.

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In the geographical range of the Spanish Ibex four perfectly
distinct areas may be distinguished, although now reduced toa
number of small isolated colonies by continued persecution from

Proc. Zoou, Soc.—1911, No. LX VI. 66

Map showing the probable former distribution (dotted areas) and present colonies (vertical lines)

of the Spanish Ibex.

966 PROF. A. CABRERA ON

the Middle Ages, when wild-goat meat was a very favoured dish
at every Spanish table. A different subspecies is found in each
of these areas, as follows :—

(a) Pyrenean area, comprising the Spanish side of the Pyrenees
and, in former times, the eastern part of the Cantabrian chain.
Tts peculiar Ibex is Capra pyrenaica pyrenaica. It may be con-
sidered as practically extinct, being today found only in the
northern extreme of the Huesca Province, about the Mount
Perdido*. Two old bucks, three females, and three or four
half-grown individuals remained there in 19077. In a recent
letter on this subject, the Count of San Juan, who spends a great
part of his time hunting in the Pyrenees, kindly informs me:
“T think that probably no more than ten or twelve Ibexes
remain in all the Pyrenean chain. A pair survived recently in
the Maladeta; somebody shot the female, and the male sought
refuge among a herd of domestic Goats and was subsequently
killed by the goatherd.”

(6) North-western, or Atlantic, area, formed by the mountains
of Galicia and Northern Portugal. The lack of suitable material
prevents correct identification of the Wild Goat found there, but
from the description by Barboza du Bocage t I surmise that it
represents a peculiar subspecies, which I do not care to describe
at present. At all events, it is well-nigh extinct, only a few
specimens, if any, remaining in the Portuguese mountains of
Gerez. Not being a Spanish race, we need not discuss it now.

(c) Central area, embracing the Sierra de Gredos and, in the
past, the ridges of El Barco, Bejar and Francia, and the hills
of Toledo. The subspecies inhabiting this area, at present reduced
to a single colony in the highest peaks of Gredos, will be named
and described below. MM. Chapman and Buck have told the
history of this Ibex so accurately that it is unnecessary to repeat
it here. The colony consists of about three hundred and fifty
head, and having been under royal protection since 1905 it is
rapidly improving.

(¢) Mediterranean area, from the mountains forming the
Guadalquivir basin, eastwards and northwards through the
sierras of the Valencia Province to the mouth of the Ebro. It
is inhabited by Capra pyrenaica hispanica (type locality, Sierra
Nevada), a subspecies not so near extinction as the Pyrenean and
Central forms. No less than six colonies, in fact, are known to
exist, the exact number of heads in each of them being unknown.
Three of these colonies are in South Spain, viz. :—one in Sierra
Nevada, another in the two parallel ridges of Sierra Bermeja and

* The Mont Perdu of the French. It is a custom with many English writers to
use French names for localities on the Spanish slope of the Pyrenees, but, in my
opinion, such a course is against commonsense. Since these localities are in Spain,
Spanish names must be preferred in every case in which there is not an English name
for them.

+ Gourdon, Bull. Soc. Se. Nat. de ’Ouest de la France, (2) viii. 1908, p. 12.

+ Mem. Acad. Se. Lisboa, 1857.

THE SPANISH IBEX. 967

Sierra de Ronda from their junction in the Sierra de Tolox, and
the third one in Sierra Morena, near Fuencaliente, under the
protection of the Marquis of Mérito. Another community of
C. p. hispanica is that of Sierra Martés, Valencia Province,
mentioned by Chapman and Buck. It is very possible that
Ibexes exist also in the mountains on the opposite bank of the
river Cabriel, a name itself derived from “cabra”; but if such is
the case, these Goats cannot be considered a different colony, since
they may swim across the river, as a specimen now preserved in
the collection of the Institute of Valencia was actually seen to do.

The two remaining colonies of this race are established on the
lower Ebro, not far from Tortosa. One of these comprises the
Sierra de Card6 and the Tivisa Mountains, where an immature
male, recently received by the Madrid Museum, was obtained.
The other colony is found on the other side of the river, on
Mount Caro.

At first glance, the three subspecies of the Spanish Ibex are
much alike, their chief differences being in the horns and in the
extent of the black markings peculiar to these Goats. The
species, as a whole, may be described as a pale brown animal
with the outer side of the limbs black, a black band on the lower
part of the flanks, and a short black mane, continued along the
back by a narrow stripe. The forehead and the beard are
blackish or very dark brown, and the belly and inner part of
the limbs white. In winter pelage there is a whitish underfur,
quite absent in summer, when the general colour is browner and
the black areas become more abruptly definite. The females lack
in all seasons the mane and the black markings of the head and
body, presenting only a blackish tint on the anterior face of the
limbs *, and it is the same with young males, in which the black
areas appear in the second or third year, becoming larger and
darker as the animal grows older. It is, therefore, very difficult
to ascertain the differences between the various subspecies when
quite adult males are not at hand.

In the typical Capra pyrenaica the dorsal line appears con-
siderably broadened on the withers, frequently forming a large
lozenge-shaped blot which in old specimens spreads laterally
over the shoulders, coming downwards to coalesce with the black
of the fore limbs. The black of the hind limbs extends upwards
over the whole external surface of the thighs and on the hind-
quarters, sometimes reaching the median stripe on the rump.
The Mediterranean C. p. hispanica has these black areas con-
siderably reduced, the dorsal stripe being not broadened anywhere,
and the black of the fore limbs reaching at most the lower part
of the shoulder and the chest, whereas on the thighs it does not
reach the haunches and is narrowed to a mere band connecting

* In one of the illustrations in Chapman and Buck’s ‘ Unexplored Spain,’ repre-
senting the shooting of Ibexes in the Sierra de Gredos, the females are erroneously
depicted with a well-marked dorsal line. In reality,this stripe is,in females and
young males, very faint or quite obsolete.

66*

PROF A, CABRERA ON
Text-fig. 195.

968

Distribution of the black areas on the body of the Pyrenean (A), Gredos (B),

and Mediterranean (C) races of the Spanish Ibex.

THE SPANISH IBEX. 969

the band of the flank with the black of the leg, thus dividing the
white of the inner side of the thigh from the general brown
colour.

The Goat of Central Spain represents a stage intermediate
between typical pyrenaica and p. hispanica by the spreading of
the black areas, this colour invading the lower half of the shoulders
and covering the whole outer side of the thighs, but not reaching
the withers nor the haunches. The dorsal stripe, as in hispanica,
is of practically the same width from neck to tail.

Text-fig. 195 clearly shows the gradual decrease in the amount
of black as the species approaches the Mediterranean coast. The
variation is to some extent parallel to that indicated by Lydekker *
for the subspecies of Capra sibirica, but in the case of the Spanish
Ibex it cannot be attributed to differences in elevation nor to
the presence or absence of snow in the localities frequented by
each race. I think it interesting to note that young males of
C. pyrenaica pyrenaica, when two years old, closely resemble,
in the black markings, adult males of the Central subspecies,
the young of the latter being in turn similar to the adult
C. p. hispanica.

As to the colour of the upper surface of the body, in winter
pelage the three subspecies are much alike, the general tint
being pale brownish grey in C. pyrenaica pyrenaica, dirty
buff, more or less clouded with black towards the lower part
of the flanks, in the Ibex of Central Spain, and a paler and less
blackened buff in C. p. hispanica. The under side of the neck
is black or dark brown in the Pyrenean and Central races, and
slightly clouded with black in the Mediterranean form. I have
never seen a specimen of true pyrenaica in summer pelage,
but Trouessart describes its colour during that season as “ gris
brun foncé.” In both the two other subspecies it is pale brown,
washed with white on the flanks. here is, however, a difference
of tint between them, the Central Ibex being browner and the
Mediterranean one redder. The colour of the Ibexes of Gredos
is near the broccoli-brown of Ridgway, whereas in C. p. hispanica
it is a tint intermediate between fawn and cinnamon. Each
hair is white at the root, after which there is an undulated space
rather curiously coloured, as it has one side white and the other
side brown. This particoloured space is followed by a broad pale
band, and the hair ends in a dark brown point. Now, the
difference in colour between the two subspecies depends on the
pale subterminal space being entirely cream-buff in the Central
Ibex, and white with a broad red ring in hispanica.

Another noteworthy difference between these two Ibexes is
found in the colour of the hind border of the thighs. In the
Central form this part is buff, abruptly contrasting with the
brown of the haunches; whereas in the [bex of the Mediterranean
region it 1s coloured like the rump and the flanks, the tint being

* P.Z.S. 1901, i. p. 91.

970 PROF. A. CABRERA ON

only a little paler. Moreover, the sides of the head, grey in the
Pyrenean and Central races, are pale buff, clouded with brownish
red, in the Mediterranean form.

Text-fig. 196.

aN 4 i
SAW

TR
ont :

a

aS
$
\

Skull and horns of adult male of the Gredos Ibex (x 1).
Madrid Museum, No. 1523.

The skulls of C. pyrenaica pyrenaica and C. p. hispanica have
been described by Forsyth Major, who gives a number of dif-
ferential characteristics, most of them, I think, either merely
individual or due to age. A reliable one appears, however, to be
found in the shape of the nasals, which are more abruptly
narrowed in front in Aispanica than in true pyrenaica. In this
respect the Ibex of Gredos is nearer to the Mediterranean race,
the border of the nasals forming an almost perfect V in their
distal third.

Writing of the horns of Ibexes from different localities, Chapman

THE SPANISH IBEX. 971

and Buck* assert that “‘ examples from the two outside extremes
(Pyrenees and Nevada) most closely assimilate in their flattened
and compressed form of horn.” If by this a lateral compression
is to be understood, my own experience bears out this statement ;
but if we must understand that the individual horn in the
Pyrenean and Mediterranean races is flatter from front to behind
than in the Gredos subspecies, on careful inspection I cannot agree
with the above-quoted authors. The only reliable method for

Text-tes NOTE

Cross-sections of left horns of the Gredos Ibex (x 4).
A. Madrid Museum. Type.
B. Madrid Museum, No. 447.
(In this and the two following figures the sections represent the horn as

being seen from the tip, the front face appearing above and the inner keel to the
right side.)

investigating the true differences, and the one I have followed
with every specimen examined, consists in the taking of a cross-
section of the horn about the middle of its length. This section
is pear-shaped, and somewhat variable even in specimens from the
same locality ; but in each subspecies it is always easily referable
to a peculiar type. Now, in the Gredos Ibex (text-fig. 197) it is

* “Unexplored Spain,’ p. 144.

972 PROF. A. CABRERA ON

invariably flatter and broader than in any one of the other sub-
species, the difference being chiefly due to the great width, in the
former, of the flat upper surface of the inner keel, between its
edge and the round part of the horn. The horn sections
of OC. p. pyrenaica and C. p. hispanica, although much alike
in their more rounded and narrow contour, also differ in form.
In true pyrenaica the portion of the upper or front face imme-
diately above the keel is markedly hollow, whereas in hispanica

Text-fig. 198.

Cross-sections of left horns of the Pyrenean Ibex (X $).

A. Mainz Museum. Type.
B. Toulouse Museum.

it is slightly convex, the Mediterranean Ibex approaching
in this the Gredos subspecies. Of course, these differences can
be appreciated only in adult males, as the horns of the young in
all the races have a somewhat rounded section with a short pro-
jecting tip corresponding to the inner keel. Every specimen
I have compared has horns with at least six annulations *.

As to the curvature and direction, the horns are absolutely alike

* The popular belief that each knot or annulation on the horns means a year in
the age of an Ibex is no more true than the one assigning the same value to the
tines of deer; but, since the number of knots depends on the horn growing, it
becomes evident that many annulations, like many tines in deer horns, always indicate
an old animal.

THE SPANISH IBEX. 973

in the three races, being twisted in a half-turn of spiral, with the
tips sometimes directed inwards and downwards, as in Capra
cylindricornis, but generally pointing upwards Thus, the horns,
when seen from the front, form a very open lyre, not unlike the
lower half of the horns of the Pir Panjal Markhor. In the females
they are very short, somewhat lyrate and quite cylindrical,
differing from those of young males in the complete absence of
keel. The size of the horns in the adult male is decidedly larger
in the Pyrenean Ibex, the two other subspecies being practically

Text-fig. 199.

Cross-sections of left horns of the Mediterranean Ibex (xX #)-

A. Madrid Museum, No. 449.
B. Madrid Museum, No. 1042.

similar in this respecs. I have neither seen nor found mentioned
any specimen from either Central Spain or the Mediterranean
area with horns about one metre in length, such as frequently
occur in Ibexes from the Pyrenees. There followsa table of horn-
measurements of adult males in the three subspecies. The
dimensions of specimens marked [Ch. & B.] after the owner's
name, are converted into millimetres from Chapman and Buck’s
‘Unexplored Spain’; those of specimens marked [W.| from
Rowland Ward’s ‘ Records of Big Game.’

PROF. A. CABRERA ON

Pyrenean Thee }

Length on

outside circum- Tip to tip. Locality. Owner.
curve. ference. |
mm. mm. mm. |
1020 260 | Valibierna. Bagneres de Luchon Museum.
970 a | | North Spain. Paris Museum.
830 be: | Bee Ordesa Valley. Bordeaux Museum.
(Sin 22 675 Ordesa Valley *. Sir Victor Brooke [Ch. & B.]. |
750 | 960 | 450 (about) + Pyrenees. Mainz Museum (Type) f.
730 | 855 585 | Ordesa Valley. Sir Victor Brooke [Ch. & B.]. |
698 | 255 495 | Pyrenees. British Museum [ W.].
630 250 | Benasque. Nantes Museum.
610 | Pyrenees. Yaris Museum.
590 | | ; Querigtiena. Toulouse Museum.
Gredos Ibex.
815° | 252 690 + Gredos. |H.M. the King of Spain.
768 | 240 595 Central Spain. / MM. Chapman & Buck [W.}.
740 | 250 530 Bohoyo. Madrid Museum (No. 447).
730 220 470 + Gredos. | Madrid Museum (No. 1523).
M5 | 245 620 Gredos. | Marquis of Torrecilla.
700 165 520 Gredos. Senor Prado Palacio.
680 245 520 Las Hoyuelas. Marquis of Viana.
650 | 260 550 Madrigal de la Vera. | Madrid Museum (No. 448).
630 260 540 Las Hoyuelas. Marquis of Viana.
613 230 | Gredos. MM. Chapman and Buck
on A [Ch. & B.]. |
Mediterranean Ibex.
850 a a | Sierra Morena. | Marquis of Mérito [Ch. & B.]. |
740 200 | 525 | Sierra Nevada. | Madrid Museum (No. 449).
730 230 585 | Sierra Nevada. | MM. Chapman and Buck
| (Ch. & B.]. |
650 Be .. | Sierra Nevada. | Bordeaux Museum.
590 220 320 South Spain. | Senor E. Cortina.
590 210 470 | Sierra Nevada. | Madrid Museum (No. 921).
580 200 Sierra Nevada. | Madrid Museum (No. 1048).
580 200 sai Sierra Nevada. | Mainz Museum.
570 250 410 Sierra Nevada. | Madrid Museum (No. 1042).
655 200 | 415

Sierra Martes.

| Senor P. Burgoyne [Ch. & B.}.

* MM. Chapman and Buck give merely “ Pyrenees”? as the locality of Sir Victor

Brooke’s specimens, but according to a letter of the owner himself, published by
Count Russell (see Gourdon, Bull. Soc. Sc. Nat. de ?Ouest de la France, viii. 1908,
pp. 6-8), they were obtained in Ordesa Valley, the Val d’Arras of the French.

+ The tip of a horn is slightly broken.

+
te

Capra pyrenaica was based by Schinz on specimens in the Mainz Museum,
which are still there, as Dr. Reichenow kindly informs me.

One of these specimens

being an adult, although not old, male in winter coat, and the winter pelage being
described first by Schinz, I think it convenient to choose it as the type.

THE SPANISH IBEX. 975

T must now proceed to the complete description of the Central
Spain Ibex. His Majesty King Alfonso XIII. has specially and
graciously permitted me, at my own request, to dedicate this
subspecies to Her Majesty Queen Victoria of Spain. I have
great pleasure in doing so, in recognition both of her love of
nature and of the fact that Royal protection only has prevented
the total extinction of this splendid ruminant.

CAPRA PYRENAICA VICTORLE, subsp. n.

Diagnosis.— An intermediate form, in size and in the extent of
the black markings, between C. p. pyrenaica and C. p. hispanica,
rather browner than hispanica in the summer coat, and with horns
similar in size to those of that race, but comparatively broader
and flatter.

Colouwr.—Adult male, summer pelage: Upper surface of body
pale broccoli-brown, sometimes inclining to raw umber, and more
or less white-washed on the sides. Neck fawn-coloured, paler, and
frequently whitish, on the under surface. A black stripe, bordered
with white hairs, starts from a large black blot covering the nape
and runs along the upper part of the neck and over the back,
reaching to the tip of the tail. On the neck the hairs of this
stripe are Jong and erect, forming a short mane. On the back
the white hairs form a narrow and not very distinct light area on
both sides of the black median line. Belly and inner side of
thighs white. A broad black band, grizzled with white at the
borders, crosses obliquely the lower part of the flanks, its anterior
point going into the white of the underparts. Thighs black,
except on the hinder border, which is light buff. The black
spreads downwards, encircling the limb above the hock and
covering the front and sides of the leg and the whole foot from
some distance above the false hoofs. The fore feet and the front
and lateral surfaces of the fore limbs are likewise black, this
colour reaching the lower half of the shoulders, the chest, and the
lower part of the ventral aspect of the neck, where it is coarsely
mixed with white. Back of the legs creamy white. Forehead
seal-brown; cheeks brownish grey; the eyes encircled with
ochraceous buff, and the muzzle and the upper lip are of the same
colour. The beard brownish black, this dark tint covering also
the sides of the lower jaw to the rim of the mouth, whereas the
middle of the lower lip is dirty white. The ears fawn-coloured on
their outer aspect, yellowish white within.

Winter pelage: The main colour of the upper parts of the body
and neck turns in winter a dirty buff, densely clouded with black
on the flanks, the hairs being white at their bases and then pale
eream-buff with a brown or blackish tip, and covering a whitish
under-fur. Throat and underside of the neck seal-brown, touched
here and there with chestnut. Black areas distributed as in the
summer coat, but not so abruptly defined, their upper borders
melting into the black clouding on the sides. The colours of the
head are practically the same as in summer, the cheeks only being
slightly paler and somewhat buffy.

976 ON THE SPANISH IBEX.

Adult female: In summer pelage the general colour is an
intermediate tint between cinnamon and fawn, paling to creamy
white on the underparts, the inner side of the limbs, and the
lateral and hinder surface of the legs. ‘The muzzle and the sides
of the face cream-buff. The fore part of the legs, from a short
distance above the knees and hocks, Vandyk-brown. Tail like
the back, with a seal-brown tip. Dorsal stripe and bands of the
flanks quite absent. In winter coat the main colour is a dark
dirty buff.

Young of both sexes, in the first year: Colour like adult females,
but somewhat paler; the markings on the legs pale chestnut.
The males begin to show the dark areas of the body in the third
year, the black tint appearing first on the chest and lower part of
the shoulders.

Skull and horns.—See above for the comparison between this
and the other subspecies.

Measurements (of type, mounted).— Length from nose to root
of tail, along the curves, 1355 mm.; tail, 130; hind foot, with
hoofs, 385; ear, 120; height at shoulder, 700 *.

Skull (of paratype, Madrid Museum, No. 1523): Total length,
264 mm.; interorbital breadth, 110 ; length of nasals along median
suture, 95; greatest width of ditto, 40; upper molar series, 68 ;
lower molar series, 75. For the horn-measurements, see the
foregoing table (p. 974).

Type.—Old male in summer coat, from Madrigal de Ja Vera, on
the southern slope of the Sierra de Gredos. Madrid Museum,
No. 448.

The differences between this subspecies and both the typical
and the Mediterranean forms of C. pyrenaica, fully discussed
above, will be, I hope, clearly shown by the accompanying figures
and plates. I have considered it unnecessary to give a coloured
figure of the Pyrenean form, as there is a tolerably good one in
Lydekker’s ‘ Wild Oxen, Sheep, and Goats,’ besides the portrait of
a young male in Cuvier’s ‘ Histoire Naturelle des Mammiféres.’
Unfortunately, the same cannot be said about C. p. hispanica.
The figure published by Schinz in ‘Monographien der Siugethiere’
is anything but good, and the one in Rosenhauer’s ‘'Thiere
Andalusiens’ is not much better, the rigidity and other defects of
the mounted specimen, after which it was evidently made, being
too faithfully reproduced by the artist. As for the Ibex of
Central Spain, in Graells’ ‘ Fauna Mastodolégica Iberica,’ there
is a plate which appears to be an attempt to represent some of
the specimens in the Madrid Museum ; but they are figured in
quite a grotesque and childish way, and the colour is entirely
false. The Ibexes on the Risco del Fraile in Chapman and Buck’s
‘Unexplored Spain’ are correctly drawn; but the illustration,
being uncoloured, cannot give a complete idea of the animal.

* In an old buck from the Sierra Nevada, in the Madrid Museum, the length
from nose to root of tail is 1190 mm.; the hind foot, 305.

SUSKehEIININstel WH IANONONGE INL: tS
‘SISNHEVaGVH ISdITTMHd VOOOGVN ¢ SISNHNVaGND ISdITIHd VOOOCVIN +

‘OULD UEUIMAN 4Se\

x Hl
FARO

AT id JI6t SAd

AS. USGL PAL ILL,

West, Newman chromo.

2. MADOQUA SWAYNEI.

1. MADOQUA PIACENTINII.

ON THE SOMALI DIK-DIKS. 977

I must not close this paper without expressing my sincere
acknowledgments to Prof. Dr. Reichenow, Mainz; Dr. A.
Ménégaux, Paris; M. A. de Montlezun, Toulouse; and M. Chaine,
Bordeaux, for helping me with measurements and other information
about specimens in their respective museums.

iy

EXPLANATION OF THE PLATES.
Prate LIT.

Capra pyrenaica hispanica.—Adult male from the Sierra Nevada, in summer coat.
Madrid Museum of Natural Science.

Prats LIT.

Capra pyrenaica victorie.—Old male (type) from Madrigal de la Vera, and adult
female trom Bohoyo, both in summer coat. Madrid Museum of Natural Science.

Prate LIV.

Capra pyrenaica victorie.—Old male from Bohoyo, in winter coat.
Madrid Museum of Natural Science.

44. On Antelopes of the Genera Madoqua and Rhynchotragus
found in Somaliland. By R. EH. Drake-Brocxmay,
MERC Selistu.Oaky., HZ.)

[Received and Read June 138, 1911.]
(Plates LV. & LVI.*)

THe DIK-DIKs.

The Somali country might justly be termed the home of the
Dik-diks,' for not only are most of the known species found there
but all the remainder at present known, save Madogua dama-
rensis, are to be found in the adjoining territories.

These little Antelopes, affording but poor sport for the big-
game hunter, have been quite overlooked by the sportsman until
the last fifteen years or so.

Dik-diks are invariably found in what might be termed the
acacia bush country, dry and arid regions where the trees seldom
grow beyond the height of bushes, and where there is sufticient
thick undergrowth to afford them protection, while at the same
time it permits of their running about freely between the bushes
and plants. In the dense aloe and sansevieria patches so
frequently met with in these parched regions, Dik-diks will nearly
always be plentiful, owing to the excellent shelter they afford for
these diminutive creatures.

Most of the small plantsand bushes supply them with food, but
the various varieties of stunted acacias are undoubtedly their
favourites. Personally I cannot recall having met with Dik-diks
in any place where these stunted acacias were not to be found.

* Wor explanation of the Plates see page 984.

978 DR. R. E. DRAKE-BROCKMAN ON

It has been doubted whether they can exist without water ;
Swayne says, talking of all the Dik-diks, that they “like to be near
water, going to drink at midday and just after nightfall,” whereas
the Somalis maintain that if a Dik-dik drinks water it will die.

In the Badminton Library, vol. 1., ‘Big Game Shooting in
East Africa,’ Mr. F. J. Jackson wrote concerning the Paa, the
local name for M/. kirkii, “it is therefore quite evident that the
juices of the vegetation on which it feeds and the dews at night
are sufficient for its requirements.”

My own experience is in entire agreement with the last-
mentioned authority, although I am not prepared to say that
Dik-diks never drink.

Those who may have observed them closely will have noticed
how a single pair will, even though disturbed, be found day after
day in much the same spot, perhaps a square acre in extent, miles
from the nearest water; here they will live all their lives if
unmolested. It is surprising to see how, after shooting the
female, the male will cling to his old haunts, living quite alone
for months together.

On one occasion I shot the female of a pair which used to live
in a small acacia grove where I frequently went to get a Francolin
for the pot ; not wanting the male I left him alone, and shortly
after left the district for seven or eight months, on the lapse of
which I returned, and while after Francolin again saw my old
friend who, not having found another mate, was living quite alone ;
there was no question about his being unaccompanied, as I saw
him every time I visited the spot. This is all the more remark-
able, as Dik-diks were plentiful enough in the locality.

There is one point I should like to emphasise before proceeding
to deal with each of the various species in turn, and that is the
presence, in a large proportion of the Dik-diks, of small white
spots on the muzzle.

Professor Linnberg has given specific rank to a Rhynchotragus
from Lake Baringo because of this peculiarity, but I venture to
think that this feature will not be found to be constant. In
specimens of M. phillips from the same district some will be
seen to possess these white muzzle-spots, while in others they are
absent.

Mapogua pHitirest Thos. (PI. LV. fig. 3.)

Phillips’ Dik-dik presents such striking variations in different
localities that for some time past I have been endeavouring to get
together a series of specimens which will show these variations
clearly and enable me to decide as nearly as possible the limits of
this interesting species.

In point of fact, were it not for the skull- measurements being so
similar, | should have been tempted to give each of the varieties
distinct specific rank, as the localities frequented by each are very
clearly defined.

Phillips’ Dik-dik was first described by Mr, Oldfield Thomas,

THE SOMALI DIK-DIKS. 979

who took as the type a specimen obtained by Mr. Lort Phillips
at Dobwein, 40 miles south of Berbera.

This spot I have been unable to locate, so can only conclude,
after carefully examining the type specimen, that it was some-
where along the Golis foothills or, perhaps, just on the top of
Mirso, which I will show is the southerly limit of the coast
or Guban variety, to which I gave subspecific rank in July 1909 *
and called MW. phillipsi gubanensis.

This subspecies is found all along the coast-belt of British
Somaliland, passing northwards into French Somaliland, certainly
as far north as Djibouti, but how much farther I have been
unable to ascertain, and eastwards towards the Mijertain country,
but how far I have not yet determined, owing to there being no
specimens available from that area.

What I propose to designate as the true I. phillipsi is the
brilliant rufous or dark cinnamon-flanked Dik. dik found through-
out the interior of British Somaliland from, roughly speaking, the
Golis Range away to the south and west into the Haud, where,
especially in the west, the animal’s flanks are so red that at
a short distance it looks rufous all over; this is in marked
contrast with the coast variety, which looks quite grey.

As one passes through the hilly country from Jig-jigga to
Harrar and also to the west and south-west of the former,
one finds another and darker Dik-dik, which in its wild state
looks of a dark red-brown colour. This is the Dik-dik to which
Mr. Neumann gave the name J/. hararensis, and so distinct does
it appear in life from J. phillips: that it fully deserves a name so
that it might not be confounded with M. phillipsi; but in my
opinion it deserves only subspecific rank, and should be called
M. phillipsi hararensis (P|. LV. fig. 2). It is to be expected that
in localities where the soils are so distinct as in the coast-belt
and the Haud one would see some variation in the coloration of
the pelage, and here in this species we have this variation very
distinctly marked; but this is not the only difference, another
equally potent factor, in the shape of altitude with its accom-
panying variations in temperature, also assists in increasing
the difference between the type species and its subspecies, by
rendering the pelage thicker.

For instance, the hair in the Guban variety is scantier and
shorter than in the Harrar and neighbourhood specimens; in
the former the individual hairs from the dorsal region of the
back measure from 13-23 mm., whereas in the latter variety they
are usually 30mm. or more. This difference renders M/. phillipsi
gubanensis a much more sleek-looking animal (EILWe ities),
The pelage of the true W. phillipsi, which is the common Dik-dik
found all over the interior of the “ Horn of Africa,” is inter-
mediate between the two.

The skull-measurements in all three are, as one would expect,
practically identical.

* Ann. & Mag. N. H. ser. 8, Vol. 4. p. 49.

980 DR. R. E. DRAKE-BROCKMAN ON

This Dik-dik is known to the Somalis by the name “ Gol ass,”
owing to its bright ved flanks. It ranges from near Djibouti in
the north and the Ennia Galla country in the west throughout
the Somali country to the east coast and as far south as probably
the 3rd parallel. A large number of the skins of this Dik-dik
are brought down to the Benadir ports for sale, from Central and
East Central Somaliland.

T might here add that I consider that little importance can be
attached to the colour of the crest, as it 1s very variable. In
some it is of a bright fulvous, in others of a dull reddish brown,
while in not a few the hairs are tipped with black. The white
eye-patch also varies in distinctness with age.

Mapboqua swayNEI Thos. (Pl. LVI. fig. 2.)

This is the smallest of the Somali Dik-diks, and is, according to
Swayne, known to the Ogaden Somalis by the name “ Guyu,” but
this name I have never heard it called myself. The word “ Guyu ”
in Somali means any living animal.

There is very little known about the exact habitat of this little
Dik-dik, owing to the fact that sportsmen have usually confounded
it with the ubiquitous MW. phillipsi. The type specimen was
bought by Swayne from a native in the town of Berbera.

IT have been on the look-out for it for some years, but have
failed to come across it either alive or dead in British Somaliland.
I first met it during my journey with the Anglo-Abyssinian
Boundary Commission in 1908, south of Ginir on the river Web,
one of the affluents of the Juba, where it was plentiful practically
all along the left bank of this river up to its junction with the
Ganale. This I took to be its westerly limit.

One of my collectors has lately brought me two specimens of a
Dik-dik from Eastern Somaliland, as far south as Obbia on the
coast ; he obtained these from a place called Gharabwein about
12 miles inland from Obbia, where they were plentiful, and
were the only Dik-diks seen, although at Eil Hur, not more than
10 miles distant, J/. phillipsi abounded and this Dik-dik
resembling JZ. swaynet was absent.

From the above I conclude that J. swaynei and the above-
mentioned species which I am about to describe stretch right
across Central Somaliland from east to west, where they are
locally distributed, and surrounded by the commoner J. phillipsi.
Neither J. swaynei nor the allied form go as far south as
Mogadishu, as my collector was unable to procure a specimen
of either species there or on the Webi Shebeleh, nor was he
able to purchase any of their skins in the market, although the
Somalis bring Dik-dik skins in thousands for sale in the coast
towns in Italian Somaliland. The only two species the skins of
which he was able to obtain were J. phillipsi and R. quentheri.

The Obbia specimens differ from those I obtained on the Web
in Western Somaliland in that there is no yellow suffusion of the

THE SOMALI DIK-DIKS, 981

grizzling on the back in the former, as there is in the latter, and
the bright rufous nose-patch is not continuous with the crest,
which may be either rufous or dull earthy brown.

In his ‘Game Animals of Africa,’ page 191, Mr. Lydekker says,
speaking of J. swaynei, “The bucks weigh but 6 lbs. and the
does even less.” Now it is an invariable rule among these small
antelopes to find the females heavier than the males. There is
usually a difference of 1 lb. between the sexes. Adult males of
Swayne’s Dik-dik weigh about 43 lbs. and the females 5 or 52 lbs.,
whereas the average weight of a buck of MW. phillipsi is 52 Ibs.
and of a doe 63 lbs. The heaviest buck of the latter species I
have ever weighed was 6? lbs., whereas the heaviest doe was
8 lbs.

I should place the range of this Dik-dik between the 5th and
9th parallels of latitude, where, although it intermingles with
M. phillips, it is not nearly so common. _ It is quite possible that
it is not to be found farther north than the 8th parallel, as
Mr. Dodds, a friend of mine, shot a large number of Dik-diks in
the Ogaden Rer Ali country, especially around Daggahbur and
Milmil, and failed to procure a single specimen of IZ. swaynei,
although both J/. phillipsi and R. guentheri were obtained.

MADOQUA PIACENTINIL, sp.n. (Pl. LV1. fig. 1.)

Although this little Dik-dik agrees, as regards its size and
skull-measurements, with J. swaynei, it differs so markedly in
coloration that I have considered it advisable to propose a new
name for it.

In M. swaynei the grizzling is very indistinct, being invariably
suffused with a dull buff or clay colour, whereas in this species
the grey grizzling is so fine and distinct that it at once attracts
attention.

The coloration of the legs in both species is similar. The
neck in M. piacentini is fairly grizzled all round, the pale throat
and buff chin patches being completely cut off from the pinkish
buff of the chest. Apart from the fine grey grizzling, the most
distinctive patches are about the head.

In IZ. piacentinu there is a bright rufous diamond-shaped nose-
patch which stands out conspicuously on the grizzled head, while
the terminal part of the long hairs of the crest in both my
specimens is of a dull creamy buff. The hairs of the crest,
however, vary so much in Dik-diks that little importance can be
attached to this feature. The ears, which are of a dark buff, have
avery distinct black edging on the outside, about 4 mm. in width
anteriorly and dwindling down to a mere edging posteriorly.
This peculiarity I have never seen in any of the other Somali
Dik-diks.

I was successful in getting only two specimens of this very
handsome Dik-dik, both males, but both present exactly similar
features. ‘They were obtained by one of my collectors at a place

Proc. Zoou. Soc.—1911, No. LX VII. 67

982 DR. R. E. DRAKE-BROCKMAN ON

called Gharabwein, within a day’s march of Obbia in the
Mijertain country, Italian Somaliland. They were inhabiting
thick aloe scrub country, and were plentiful im a locality some
little distance from water; they were very local and appeared to
be surrounded by JZ. phillipst.

The following are the measurements in the flesh of the type
specimen—a male in the British Museum, No. 356.

Hieadwandgbodiyaecca-ue oboe crcese aes A475 mm.
Wail cops ene aire Pd are tee Tien cae cae SS
Himd foGbiweamaiasithesacchcael apie. yen
A OE eee eer eee Sr omOGnE On: San ¢ suena CA He
IW ielig iG Terenas ieeunce Greene Tasers 2 lbs
Skull-measurements.
WiNeiseyll lWenaventlin " Qaqeeneusscncnesosoaconscce 92 mm.
Teiniseul Weraverslay 843 occescnaccgac ooacecacoece SOi
Posterior edge of orbit to gnathion 609 ,,
iWippericheek-teebtht..-.sen.c-cedeeas 4 Ser oS
IDeineid a Oi Weslo q- che scecone ococasseocc NGO
iBreaditiwotenasal sehen eee saa 8:9),

In comparing the above skull-measurements with a typical
M. swaynei, it will be found that they are practically identical.

This bright and beautiful Dik-dik I propose to name in honour
of my friend Mr. Piacentini, the Acting Consul-General for Italy
in Aden, through whose kindness and help I was able to send my
collector to the Mijertain country.

RHYNCHOTRAGUS GUENTHERI Thos.

This Dik-dik, known to the Ogaden Somalis by the name
‘“¢Ghussleh” or ‘‘ Gussuleh,” owing to its habit, when startled, of
dashing off in leaps and bounds, giving vent to a peculiar
whistling ery which sounds like “ Ghuss-Ghuss-Ghuss,” is chiefly
found in Western and West Central Somaliland. Swayne tells us
that he first met with it when travelling in a south-westerly
direction in the Rer Amaden country, 7. e., roughly speaking,
between the 7th and 8th parallels of north latitude; this is its
northerly limit, as Mr. J. H. Dodds tells me that during a recent
journey he made in Western Somaliland he never met this Dik-dik
until he got to Daggahbur, a well-known Ogaden watering-place
near Milmil.

Starting from Daggahbur in the north this Dik-dik passes in a
southerly and south-westerly direction for Dolo at the junction of
the Dawa and Ganale rivers, and then crossing the Juba is
ubiquitous throughout north-eastern British East Africa. I do
not think it reaches the coast anywhere, being there replaced by
R. kirkii. In Somaliland proper its easterly and south-easterly
limits are still unknown.

It probably extends for a considerable way down the Webi

THE SOMALI DIK-DIKS. 983

‘Shebeleh river, very nearly reaching to the coast in the Hawiya
country, as thousands of their skins are yearly sold in the market
in Mogadishu by the Hawiya Somalis. Theyare either caught in
native traps or shot with bow and arrow.

RHYNCHOTRAGUS KIRKIL Ginth.

Kirk’s Dik-dik, which was named so far back as 1880, inhabits
only the most southern angle of the Somali country east of the
Juba River; it, however, extends across that river and south-
wards into British East Africa as far as Kilimanjaro. How far
north it extends into Somaliland proper Lam unable to say, as the
material at present available is insufticient.

The type specimen came from Brava on the Benadir coast of
Italian Somaliland, but my collector failed to get me any specimens
from Mogadishu, so [ conclude that it does not extend farther
north than this, being replaced on the coast by JZ. phillipsi and
in Central Somaliland by 2. guentheri, of which hundreds of skins
are offered for sale in the market in Mogadishu. I have only
been able so far to procure the skins without head-skins or skulls
from that locality, so am unable to definitely state whether the
skins belong to A. guentheri or an allied form; they certainly
appear to agree with my specimens of the former.

RHYNCOTRAGUS CORDEAUXI Dr.-Br.

This Dik-dik, which was named by me last year, should really
be included among the Abyssinian Dik-diks, together with
f.erlangert, M. phillipst hararensis, and Rk. guentheri wroughtoni,
but like I. phillipsi hararensis, it will probably be found to
inhabit the north-eastern part of the Esa country.

It was first obtained by me in the bush country to the west and
north-west of Dirre Dawa, and is at present, so far as I am aware,
recorded only from the Danakil country:

I traced it as far west as the Gurgurra River, one of the
tributaries of the Hawash, and it is this latter river that marks
out its westerly limit; the northern edge of the Harrar and
Arussi plateaux mark its southern limits, but how far north
and north-east it strays I am at present unable to definitely
‘state.

This is a fine species, and most resembles in appearance
M. phillipst gubanensis, only it is a much bigger animal.

RHYNCOTRAGUS ERLANGERI Neum.

Erlanger’s Dik-dik hails from Eastern Abyssinia. As one
descends from the great Arussi plateau on the low-lying country,
towards the east, called by the Gallas ‘“‘ Gamogi,” this is the
Dik-dik one finds.
If a line be drawn on the map between Harrar in the north
-and Ginir in the south, this line will cut through an acacia-bush
67*

984 ON THE SOMALI DIK-DIKS.

country, more or less intersected by rivers and streams which drain
the great Arussi plateau; this, the Ennia Galla country, is where.
Erlanger’s Dik-dik is found.

The type specimen was obtained at Sheikh Hussein, which is.
30 or 40 miles or so east of the edge of the plateau, while my own
specimen was obtained some 20 miles to the west of Sheikh
Hussein opposite the foothills of Mt. Abu el Kassim, on the south
-bank of the river Wabi.

So far as I am aware, these are the only two specimens recorded,
so that at present its limits must remain undefined ; but I fancy I
shall be fairly accurate in suggesting that its habitat probably
lies to the west of the Harrar-Ginir line, being replaced to the east:
of it by WZ. phillipsi hararensis and to the south by IZ. swaynei.

RHYNCHOTRAGUS GUENTHERI WROUGHTONI Dr.-Br.

I have now come to a very interesting subspecies, namely
R. quentheri wroughtoni. The only specimen recorded is the type,
which I obtained on the north bank of the Wabi River among the.
foothills of Mt. Abu el Kassim.

The presence of this subspecies in a spot so far removed from
the natural habitat of the species can only be explained by its.
having at some period or other found its way up the river Wabi,
which is one of the main tributaries of the Webi Shebeleh, which
flows through the vast area inhabited by &. guentheri. Its
darker coloration and large ears are probably accounted for by its
environment, namely the dense acacia bush on the banks of the
river; the size and shape of its ears certainly point to this.

I am inclined to think that this subspecies will only be found
close to the river, as both MW. erlangeri and R. swaynei are to be
found within a short distance of it.

EXPLANATION OF THE PLATES.

Puate LV.
Fig. 1. Madoqua phillipsi gubanensis. | Fig. 2. M. phillipsi hararensis.
Fig. 3. Madoqua phillipsi.

Prate LVI.

Fig. 1. Madoqua piacentinii. | Fig. 2. Madoqua swaynei.

t=}

ON WING-CLAWS IN YOUNG BLACK-BACKED PORPHYRIOS. 985

EXHIBITIONS AND NOTICES.
June 27, 1911.

FREDERICK GILLETT, Esq., Vice-President,
in the Chair.

The Secretary read the following report on the additions made
to the Society’s Menagerie during the month of May 1911 :—

The registered additions to the Society’s Menagerie during the
month of May were 445 in number. Of these 163 were acquired
by presentation, 200 by purchase, 41 were received on deposit,
25 in exchange, and 16 were born in the Gardens.

The total number of departures during the month, by death
and by removals, was 204.

Amongst the additions special attention may be called to the
following :—

1 African Rhinoceros (2hinoceros bicornis) ¢, from Nairobi,
received from R. B. Woosnam, Esq., C.M.Z.S., for H.M. THe
Kine’s African Collection, on May 19th.

1 Californian Sea-Lion (Otaria californiana) 2, from the
North Pacific Ocean, purchased on May 10th.

2 Three-coloured Parrot-Finches (Lrythrura trichroa), new to
the Collection, from New Guinea, received in exchange on
May 4th.

A Collection of 38 Fishes, received on May Ist, comprising the
following species, all new to the Collection :—

4 Sword-tails (Xiphophorus helleri) from Mexico, 2 Fighting Fish
(Betta splendens)from Singapore, 2 Fan-tailed Cyprinodons (ivulus
flabellicauda) from Mexico, 2 Elegant Cyprinodons (Haplochilus
elegans) from the Niger River, 2 Chaper’s Cyprinodons (Haplo-
chilus chapert) from Sierra Leone, 2 Timid Cyprinodons (Haplo-
chilus panchax) from Cochin India, presented by P. Arnold, Esq. ;
2 Poey’s Cyprinodons (Rivulus poeyi) from Para, 6 Ocellated
Cyprinodons (fivulus ocellatus) from Santos, 10 Zebra Fish
(Dania rero) from Bengal, presented by G. A. Boulenger, Esq.,
F.Z.S.; 2 Freshwater Flying-Fish (Pantodon buchholzi) and 2
Gular Cyprinodons (fundulus gularis) from the Niger River, and
2 Rainbow Fish (Zrichogaster labius) from Bengal, purchased.

My. D. Seru-Smiru, F.Z.8., the Society’s Curator of Birds, ex-
hibited two immature Black-backed Porphyrios (Porphyrio melano-
notus) which had been bred in the Gardens, and remarked upon
their possession of a well-developed claw (text-fig. 200, p.986) onthe
pollex. Although these wing-claws were said to be functional
only in the Hoatzin amongst living birds, the exhibitor believed
that they were so also in the present species and also probably in

986 DR. W. T. CALMAN ON THE BRINE SHRIMP.

‘the Common Moorhen, these birds using them in climbing
amongst reeds and herbage.

Text-fig. 200.

Wing of young Porphyrio melanonotus with well-developed claw.

Mr. J. Lewis Bonuorse, M.A., F.Z.S., exhibited a pair of
Egyptian Desert-Mice (Weriones crassus) which showed a darker
and more rufous colour than normal examples. This coloration
had been artificially produced by keeping the animals in a moist
atmosphere at 80° Fahr. They were first exposed to these con-
ditions on the 7th of April, and a month later were conspicuously
darker ; after that the darkening process still continued, but more
slowly, and they appeared for some time previous to being killed
to have reached a limit to their darkening. During this same
period, and owing to the fine weather, other examples had been in
a temperature that rose during the day to 90°, falling at night to
60° or even lower. This had apparently produced no change in
their coloration. The change in the examples exhibited was:
therefore probably due rather to the humidity than to the temper-
ature of the atmosphere. Mr. Bonhote was therefore inclined to
think that the pale colour of desert animals was due to the
extreme dryness of the atmosphere rather than to any special
assimilation of their colour to the surroundings.

Dr. W. T. Catman, F.Z.S., exhibited a number of living
specimens of the Brine Shrimp (Artemia salina), bred from
Tidman’s Sea Salt. The Brine Shrimp, a small Crustacean

THE SECRETARY ON THE AFRICAN RHINOCEROS. 987

belonging to the Sub-class Branchiopoda, was found in various
parts of the world, living in salt lakes and in the shallow
ponds in which sea-water is exposed to evaporation for the
manufacture of salt. It formerly occurred in England, but had
probably long been extinct in this country. An accidental
observation recently made at the Natural History Museum
showed, however, that it wasa very easy matter to obtain a supply
of living specimens. ‘“'Tidman’s Sea Salt,” as sold in the shops,
frequently, if not always, contained living eggs of Artemia, and
an 8 °/, solution, allowed to stand for a few days, was found to
contain a swarm of nauplius larve. The first attempt at rearing
these failed owing probably to lack of food-material in the water.
The juice of green leaves pounded in a mortar and strained
through muslin was found to be a suitable food, and the addition
of a few drops of this at intervals of about a week enabled the
specimens exhibited to be raised to maturity. All of them were
females, and swarms of larve of the second (parthenogenetic),
generation had appeared.

The Secrerary remarked that on a recent visit to the Ostrich
Farm of Mr. Carl Hagenbeck at Stellingen, near Hamburg, he
had seen in the incubator fertile eggs of Struthio massaicus from
German East Africa, S. australis from South Africa, and S. mo-
lybdophanes from Somaliland, the eggs all having been laid at
Stellingen. A. Reichenow (‘ Die Vogel Afrikas,’ vol. i. p. 7) had
already described and figured certain specific differences in the
number and arrangement of the pits on the eggs of these species.
He himself had been interested to notice that the eggs of the
Masai Ostrich were much larger than those of the others, more
spherical in shape, and very smooth and porcelanous in texture.
Those of the Cape Ostrich were somewhat similar in shape and
texture, but were much smaller; Mr. Hagenbeck had informed
him that a pair of the Masai Ostrich bred by himself and sent out
to the Cape were regarded by expert ostrich farmers there as
unusually large birds. The eggs of the Somali Ostrich were
larger than those of the Cape Ostrich, but smaller than those of
the Masai species, and were markedly oval in shape witha mor gleee
less polished surface.

The SecrETARY also remarked that on his recent visit to
Mr. Hagenbeck’s Zoological Park at Stellingen, near Hamburg, he
had the pleasure of seeing a fine young pair of the common African
Rhinoceros, obtained from British East Africa, the exact locality
being unknown. The male closely resembled the ordinary figures
and mounted examples of the species, in that the skin appeared
to be smoothly stretched over the sides of the body, but the ears
were fringed with long tufts of hair. The female, on the other
hand, had no hair on the margin of the ears, and the general
external appearance was very different. At first sight it seemed

988 MR. R. I. POCOCK ON HYBRID FOALS,

as 1f it were in very poor condition, the ribs standing out through
the skin, but closer inspection showed that in reality the skin
of the flanks was disposed in thick, permanent folds, arranged
roughly like ribs. Thinking it possible that these differences
might indicate the existence of distinct races of the Rhinoceros,
on returning to London he had at once examined the Society’s
own pair of examples of this species, both of which had come from
British East Africa, probably somewhere near Nairobi. The
female, purchased in 1906, had the ears unfringed with hair, like
those of Mr. Hagenbeck’s female, but the rib-folds on the skin
were no more than indicated, although there were very heavy
permanent folds round the neck. In the male, obtained in the
current year from Nairobi as part of the King’s African Collec-
tion, the ears were fringed with hair as in Mr. Hagenbeck’s male,
whilst the rib-like folds on the skin were extremelystrongly marked,
as in the case of Mr. Hagenbeck’s female. The presence or
absence of the marginal fringe on the ears was therefore probably
either an individual or a sexual character. In the absence of
knowledge of the exact provenance of all the four examples,
nothing could be said as to whether or no the presence of the rib-
like permanent folds on the body were racial. Their existence,
however, as well as the presence of the heavy fold round the
neck, showed that it was not correct to distinguish the Asiatic
Rhinoceroses from those of Africa by the presence in the former
of permanent skin-folds. The neck-fold was almost identical ir
both, whilst, although they were differently arranged, deep body~
folds occurred in both.

Mr. R. I. Pocock, F.R.S., F.Z.S., Superintendent of the
Society’s Gardens, exhibited a photograph (text-fig. 201) of a
foal born in the Gardens on June 21st and bred between a male
Somaliland Wild Ass (Zquus asinus somaliensis) and a female
Mountain Zebra (Hquus zebra) and made the following remarks :—

“The period of gestation, dating from the day of service to the
birth, was 12? months. The general colour of the foal is sandy
fawn, the ground tint of the legs being markedly whiter. The
ears are long, as in both parents, and have a large apical black
patch, fading inferiorly in front to brown, and a brown transverse
basal stripe, running upwards mesially: a corresponding basal
stripe is present on the ear in the dam but not in the sire.
There is no white tip to the ear such as is seen in all Zebras.
The lips and area round the nostrils are black, and there is no
white on the muzzle, such as is seen in all typical Asses. Half-
way between the forehead and the muzzle there is an area
covered with many close-set narrow brown stripes and some very
faint stripes are traceable on the lower edges of the under jaw.
The mane is like that of the sire (text-fig. 202), black in the centre
and sandy fawn externally, the pale external portion showing no
trace of breaking up into evenly spaced tufts, such as are seen in

MR. R. I. POCOCK ON HYBRID FOALS. 989

Text-fig. 201.

ee

er on

i

Text-fig. 202.

Somaliland Ass, the sire of the hybrid foals.

990 MR. R. I. POCOCK ON HYBRID FOALS.

all Zebras. A narrow black spinal stripe extends from the mane
to the black tip of the tail, but it is indistinct over the hind
quarters and on the upper side of the tail where the hair is long,
and recalls the dorsal and caudal mane or crest seen in the foal
of Grévy’s Zebra. There area very distinct black shoulder-stripe
and a few abbreviated stripes both in front and behind it; there
are also indistinct traces of close-set stripes on the lower border
of the neck and a deep black belly stripe; but for the rest the
body is unstriped and of a tolerably uniform sandy fawn colour
all over, like that of the sire. Both front and hind legs are
marked with strong black stripes, broader and more numerous.
than in the sire but much less numerous and more widely spaced
than in the dam. On the inside of the legs they extend just
above the knees (carpus) and hocks (tarsus), but externally those
of the hind leg reach almost to the stifle-joint (Anee), while those
on the front leg reach to about the same height. The callosities.
on the front leg are of medium size, being much smaller relatively
than in the dam, but actually of about the same size as in the
sire; and, as in the latter, there is no dewlap and the hairs along
the spine project backwards.

In general appearance this foal, which is a female, decidedly
favours the sire on account of the absence of stripes on the body
and the sandy fawn ground-colour. But it may be noted that
in the presence of the spinal stripe, the shoulder-stripe, and the
stripe on the base of the ear, it shows much greater similarity to
the typical form of African Ass, as exemplified by domestic
breeds, than to the race to which its sire belongs.

Whether the stripes will become more numerous as age
advances, remains to be seen.

This appears to be the first record of the birth of a hybrid
between the Somaliland Ass and the Mountain Zebra. Several
crosses between the domestic Ass and this Zebra have, however,
been described. The best extant account was given by F. Cuvier
(Hist. Nat. Mamm. iii. pl. 315, 1824), who accurately described
and figured a hybrid produced by a male black Spanish Ass and
a female Mountain Zebra. This animal agreed very closely with
the one just born in the Gardens, except that the ground colour,
when the animal was fourteen years old, was dark grey even on
the legs and there were distinct spots on the basal half of the tail.
The distribution of stripes was practically the same in the two;
and Cuvier’s figure shows no dewlap on the throat. The
extension of a crest of hair along the spine from the mane to the
tail and the presence of the basal stripe on the ear were noticed by
this author. The difference between this specimen and the one
born in the Gardens in colour of the body and legs is probably to
be explained by the blackness of the coat of the sire; but
St. Hilaire, who saw the foal when newly born, mentioned that
its general colour was yellowish chestnut ; at two years, however,
it was grey and this tint was retained until death. The period

MR. R. I. POCOCK ON HYBRID FOALS. 99}

of gestation was a fortnight over twelve months, being shorter by
one week than in the case of the animal born in our Gardens.

Another hybrid of the same kind is figured on plate 28 of the
volume on Horses in Jardine’s ‘ Naturalist’s Library.’ The
animal seems to have been much more copiously striped than our
specimen. Many strong but abbreviated stripes are shown running
along the saddle behind the withers; the leg stripes extend
farther up the quarters and the body, head and neck are marked
with indistinct wavy and close-set stripes. The croup, however,
seems to have been self-coloured sandy fawn like the rest of the
body. In this case it is not known which of the two species was
sire and which dam.

In the ‘ Knowsley Menagerie, p. 73, two hybrids between
Mountain Zebra mares and Asses of African descent are
described. One sired by a Maltese Ass is represented by the
right figure of the pl. lvii. The ground-colour of the body is
dark grey, that of the belly and legs white; the face below the
eyes is tan, and there is no white on the muzzle; the ears are
large with the tip broadly black and a broad stripe near the base ;
the mane is grey and unstriped. There are no stripes on the
face; but the body and neck are covered with narrow wavy
stripes which break up into small spots upon the hind quarters ;
the shoulder-stripe is very distinct, broader than the others and
forked; the belly is unstriped, but the legs are distinctly striped.
Except for the presence of spots on the croup, this animal is rather
like those figured in the ‘ Naturalist’s Library.’

The second specimen, sired by an ass of unspecified breed, is:
described as grey with an indistinct cross and a few narrow dark
stripes on the shoulder and fore legs [nothing is said about the
hind legs]; the upper side of the tail, which is elongate and
tufted, is stated to be slightly banded ; and the ears are said to
be moderate. Attention is drawn to the presence of scarcely any
stripes on this animal, which is contrasted on that account with
the one sired by the Maltese Ass. Although according to the text:
and the legends of the plates, this animal is unfigured, the
description applies very closely to the specimen represented by
the left-hand figure of pl. lvii., which purports to be a hybrid
between a male Hemione (=Onager) and a female Zebra.

This figure represents an animal sandy fawn in colour with
the legs striped, but somewhat sparsely, only slightly higher than
the level of the belly which is lighter than the flanks; the head
is fawn with some narrow close-set rufous stripes in the middle
about half-way between the eyes and the nostrils; the muzzle is
ashy grey, without any white; the ears are moderately long, with
a black tip; the mane is black in the middle, white externally,
the white hairs showing a decided tendency to break up into tufts
as in all zebras and quaggas; continuous with the black mane is.
a black spinal stripe; there is a distinct black shoulder-stripe,
followed by several abbreviated and less distinct stripes upon and

992 MR. R. I, POCOCK ON HYBRID FOALS.

just behind the withers. In the text this animal is merely
described as having the shoulders and legs banded. Its re-
semblance to the hybrid just born in the Gardens is very close
and, indeed, surprising considering the difference in coloration
and general character between the Onager and the Wild Ass of
Somaliland. Jin our hybrid, however, the ears are decidedly
longer and the mane shows no signs of being striped. Apart from
these differences, there is little to choose between the two. But
these differences are sufficient to make one hesitate in adopting
the suggestion that the animal described as a hybrid between an
Onager and a Mountain Zebra may have been a hybrid between
a Domestic Ass and a Mountain Zebra.”

[Supplementary note added July 4th, 1911.]

Since the above given account was read a hybrid foal (text-
fig. 203) between the same Somaliland Wild Ass and a Chapman’s
Quagga (1. quagga chapmanni) has been born. The period of
gestation was twelve months and four days from the date of

Text-fig. 203.

Chapman’s Quagga and hybrid foal.

service. The foal is very like the Mountain Zebra hybrid. The
ears, however, are smaller, as was to be expected from the
relatively smaller ears of the Quagga dam. ‘The ground colour of
the body, too, is a little paler, while the legs are only slightly
paler than the body. The leg stripes are less distinct and less
numerous and the insides of the legs are scarcely banded. The

MR. R. I. POCOCK ON HYBRID FOALS. 993

shoulder-stripe is shorter and simpler ; the spinal strip is black and
zig-zag over the saddle, and very faint, short, narrow, close-set:
stripes are detectable on each side of it along the back and
elsewhere on the body, the hairs of these ‘ ghost-stripes’ being a
shade darker and glossier than the spaces between them. The
spinal crest is not so pronounced; the apical black patch on the
ear is smaller, and the basal stripe, although much narrower,
is decidedly blacker. Faint narrow stripes are present on the nose
as in the first described hybrid; the lips and area round the
nostrils are black, and the two patches above the nostrils which
are dark tan in the dam are dark greyish brown in the foal,
and somewhat sharply contrasted with the narrow lighter area
that intervenes between them and the general sandy grey hue
of the nose. ;

The left-hand figure of pl. lviii. of the ‘ Knowsley Menagerie’
represents a hybrid between a Domestic Ass and a Burchell’s
Quagga (1. quagga burchelli). This hybrid appears to differ from
the one above described in having a few quite distinct brown
stripes on the body and very few stripes on the legs. The
ground colour of the legs, moreover, is markedly whiter. Since
Burchell’s Quagga differs from Chapman’s in having the legs
whiter and almost stripeless, the difference in the coloration of
the limbs between the two hybrids is not a matter for surprise.

In connection with the two hybrids born in the Gardens and
those that have been bred elsewhere previously between the same
species, namely H. asinus and L. zebra, or LH. quagga, irrespective
of the exact race or breed of the species, the following points
may be noticed. The white muzzle of H. asinus is eliminated.
In other respects asinine characters are dominant over zebrine
and quaggine characters, as is shown by the absence of stripes
on the mane, the disappearance of the white tip to the ear, and the
suppression, partial or complete, of the stripes on the neck, head,
body, and quarters. Even when stripes are visible on these areas
they do not resemble in width and arrangement those of Mountain
Zebras and Quaggas, but are more suggestive of the narrower
stripes of Grévy’s Zebra (L. grevyi), as in the case of some horse-
quagga hybrids bred by Prof. Cossar Ewart. Examples of typical
#H. asinus carry a dark basal patch on the ear, a spinal and
a shoulder stripe, and very frequently distinct or indistinct bars
on the legs. The spinal stripe is also frequent on all Asiatic asses;
and the Mongolian species (4. hemionus hemionus) sometimes
shows traces of a shoulder-stripe and of leg-stripes on the knees
and hocks as well. The Tibetan Wild Ass (Z. kiang) has a large
dark basal patch on the ear, and spinal, shoulder and leg stripes
are commonly visible in many horses. Although the basal ear-
patch, the shoulder stripe, and spinal stripe are absent in typical
examples of H. asinus somaliensis, no one doubts that this race
is descended from asses bearing the marks in question. Inter-
mediate forms indeed, with very narrow spinal and shoulder stripes
and a dusky patch on the ear, connect the Somaliland Ass with

994 MR, F. BE. BEDDARD ON

ordinary domestic varieties; and in all the many foals born
in the Gardens between our Somaliland Ass and domestic asses of
English and Spanish breeds, the ear-patch, shoulder and spinal
stripes were present as in the dams.

The above stated facts suggest that, with the possible exception
of the ear-patch in horses, the shoulder and spinal stripes as well
as the stripes on the legs have been lost comparatively recently
by the species that are without them.

PAPERS.

45, Contributions to the Anatomy and Systematic Arrange-
ment of the Cestoidea. By Frank HE. Bepparp, M.A.,
F.R.S., F.Z.S., Prosector to the Society.

[Received May 28, 1911: Read June 27, 1911.]
(Text-figures 204-215.)

Il. On two New GENERA OF CESTODES FROM MAMMALS.

The following communication to the Society contains an
account of the anatomy of two species of Tapeworms, contained
in the collection of Cestoidea belonging to the Society. I
feel it necessary to refer them to new genera, and the facts which
I shall presently bring forward will I hope justify my so doing.
The first of these species is certainly near to Zhysanosoma, and is,
as I think, clearly to be referred to the same genus as a species
recently described by me * and temporarily reterred to the genus
Thysanosoma. I pointed out, however, in that paper a number
of differences from other tapeworms referred to the genus
Thysanosoma, and intimated that it probably would be necessary
to institute a new genus for the worm. ‘This expression of
opinion is justified by the discovery of a second species which
is clearly of the same genus. The second species considered in
the present paper was placed by me temporarily, and in a report to
the Prosectorial Committee, in the genus Sertiella. It shows,
however, some aitinities to Tenia (sensu stricto) and seems on
account of various peculiarities, which in due course will be
pointed out, to demand the creation of a separate genus.

T shall commence with an anatomical description of the new
species allied to Thysanosoma; this I place in a new genus which
I propose to call Thysanotenia.

Thysanotenia lemuris, gen. et sp. n.

An example of a Black-headed Lemur (Lemur macaco), which
had lived in the Gardens three years and three months, contained

* See No. I. of this series, P. Z.S. 1911, p. 651.

NEW MAMMALIAN TAPEWORMS. 995

two perfect examples, and a number of large fragments, of a
tapeworm which I refer to the above new genus and species.
There is, as it appears to me, no doubt whatever that these
tapeworms belong to the same genus as a species which I have
recently described in the “ Proceedings” * of this Society as Thy-
sanosoma gambianum from the Gambian Pouched Rat (Cricetomys
gambianus). ‘The external characters are in very near agreement.
The present species agrees with Zhysanosoma gambianum in the
unarmed head, in the fact that the segments increase in length
posteriorly but are never longer than broad, that the genital
apertures are unilateral, and that the eggs are enclosed in a
numerous series of paruterine organs in the fully mature
segments. This set of characters is sufficient to refer the present
species, as well as Thysanosoma gambianum, to the subfamily
Thysanosomine of the Anoplocephalide.

Text-fig. 204.

Thysanotenia lemuris, about twice natural size.

Of the two specimens which possessed a scolex, the larger
(text-fig. 205) measured at least four inches in length, and was
probably longer as it ended abruptly and without a definite termina]

* Above, p. 651.

996 MR. F. E. BEDDARD ON

proglottid, such as was visible in the younger, smaller, example (text-
fig. 204). Furthermore, among the fragments was one measuring
nearly five inches in length, which may, for what I know to the
contrary, have belonged to the first mentioned individual, since it.
consisted entirely of mature proglottids, which were of greater
length individually than the terminal proglottids of the specimen

Text-fig. 205.

Thysanotenia lemuris, a second specimen. Above and to the left is the scolex
more highly magnified; below and to the right, two segments of another
individual showing the papillae which bear the genital pores.

described and figured in this communication. The longest
proglottids of the mature fragment were rather more than 3 mm.
long, and rather less than 3 em. wide at the posterior end of the
proglottid, which is considerably wider than the anterior end.

The proglottids have no fringes posteriorly such as are
present in Thysanosoma and provide the reason for the name
given.

NEW MAMMALIAN TAPEWORMS. 997

The scolex (text-fig. 205) is of moderate dimensions, not very
minute, and quite visible as such by the naked eye. It is
more or less spherical, and is marked by some black pigment
below the level of the suckers. A pigmentation of the
suckers is not uncommon among the Cestoidea, and it seems
occasionally to pervade the scolex also. The suckers have a
circular orifice, and present no marked features on examination
with a lens. ‘They lie some way below the summit of the scolex,
and are directed laterally ; there is no trace, that I could see, of
an upward direction. The smoothly rounded upper end of the
scolex shows no hooks. ‘The suckers lie near to the lower end of
the head, which is followed by a distinct neck of 2 or 3 mm. in
leneth, in which the strobilisation was not apparent. The
strobila are at first much broader than long, and each overlaps
its successor at the edges. In the larger specimen, the dimensions
of which are given above, the proglottids did not attain to an
equality in length and breadth until about three inches or so
from the scolex. ‘They never attain to a length which is greatly
in excess of their breadth, as will be apparent from the measure-
ments which I have given above. It is noteworthy that in the
smaller of the two individuals which are described here, the very
few last segments alone showed a length equal to their breadth.
In this worm (text-fig. 204) the last proglottid had an oval form,
and was of less diameter than those which immediately preceded
it. This form is usual for the last proglottid, and serves to
emphasize the fact that this specimen was complete and had
not begun to cast off proglottids. The ripe proglottids differed
from the anterior ones in their greater thickness, and in the fact
that their surface was marked by a longitudinal furrowing,
caused as I should imagine by unequal distention with embryos.
The shape of these proglottids also was different from that of
those lying in front, in that they were hourglass-shaped, with,
however, as has already been mentioned, a greater diameter
posteriorly than anteriorly. The genital pores are unilateral
without exception from end to end of the body. They are not at
all visible until the segments have begun to increase in length,
and are perhaps less obvious upon the very distended posterior
proglottids. The pores are particularly obvious in this species,
on account of the fact that they are borne upon slender processes
of the Body-wall, as will be more specially described later in
connection with the generative organs. These genital processes
lie near to the anterior border of the segments.

If the above account of the external characters of this tape-
worm be compared with my account of the external characters of
Thysanotenra gambiana, it will be seen that the two species,
though agreeing in many features, nevertheless show several
points of distinction, which enable them to be fully defined
as distinct species from these external characters only. It is
particularly to be noted that the mdividual of the two examples
of Thysanctenia lemuris which I regard as a not fully developed

Proc. Zoou. Soc.—1911, No. LX VIII. 68

998 MR. F, E. BEDDARD ON

worm is like the adult Thysanotenia gambiana in its characters.
Thus the proglottids do not increase at all appreciably in length
until the very end of the body, those anterior to this being
broader than long. It is, however, clearly a difference between
the two species that there are in Thysanotenia lemwris a much
longer series of proglottids of greater length than diameter, and
that the conical projection of the body which bears the generative
pore is more pronounced. Furthermore, while the species described
in the present communication has along neck, there is no neck in
Th. gambiana, the strobilisation commencing immediately after
the scolex. Finally, 7h. lemuris has black pigment in the scolex,
which is wanting in Zh. gambiana. There are thus several
obvious differences between the species, which as I think prevent
any confusion between them.

In transverse sections (text-figs. 206, 207) it was only ‘possible
to see a single water-vascular tube on each side. This is of con-
siderable diameter, though it fluctuates in size from place to place

Text-fig. 206.

Thysanotenia lemuris. Two sections across a proglottid, one of whicn (on the
right) shows the genital papilla. .The half only of each complete section is
shown. R&.S, Receptaculum seminis ; Sy.d., sperm-duct ; Z., testes scattered in
parenchyma; Va., vagina; W., excretory tube.

and is connected with its fellow in each proglottid by a transverse
tube, which hes near to the posterior boundary of the proglottid.
The single water-vascular tube of this species appears to represent
by its position the innermost of the two tubes found in 7hysano-
twnia yambiana. I could find no network of minute tubules arising
from these, such as are obvious in the last mentioned species.
Genital organs.—On the whole, the genital system of the species
dealt with in the present communication is not very like that of
Thysanotena gambiana. ‘There is, however, a very important

NEW MAMMALIAN TAPEWORMS. 999

point of agreement, in the similarity of the enclosure of the eggs
in numerous paruterine organs, which perhaps outweighs the
various dissimilarities which will be dealt with in considering the
testes and the ovaries and their ducts. The testes occupy in
Thysanotenia lemuris a quite different position in the body from
that which they occupy in Zh. gambiana. They lie in the former
species entirely between the water-vascular tubes, and in the
medullary region of course of each proglottid. Like the other
organs of the genital system, they commence to be visible rather
late in the body, thus contrasting very markedly with the
conditions obtaining in the second genus described in the present
paper. The testes lie mainly in the posterior region of each
proglottid, but in front of the transverse water-vascular vessel.
They form continuous rows each only one deep, and not, except
perhaps here and there, at all crowded. The testes are also
extended anteriorly to quite the front end of the proglottid on
either side of the ovaries.

The cirrus sae is divided, but not sharply, into two regions.
The terminal part which encloses the cirrus is narrow, and this
tube expands posteriorly into an oval vesicle of much greater
dimensions. ‘The whole structure has in fact much the shape of a
soda-water bottle. The hinder part of the sac is about on a level
with the receptaculum seminis and the sac itself is gorged with
sperm, the whole of the available space being filled with a mass of
sperm. It is encircled by a thick layer of muscle fibres which
run longitudinally to the longer axis of the sac, and form a
continuous coat passing in this direction to all round the sac
when it is viewed in longitudinal section, that is in transverse
sections of the proglottid. At the posterior end the thick
muscular layer is interrupted for a minute space, to permit of
the entrance of the sperm-duct which narrows greatly at its
entrance into the cirrus sac, or rather into that part of the
cirrus sac which is specialised as a vesicula seminalis. This
narrow region of the vas deferens immediately widens out
into a long tract of duct, which may be regarded as a second
vesicula seminalis. This tube is wide, quite as wide as in species
of Lertiella, and is coiled upon itself as it passes back towards
the opposite extremity of the body. This region of the sperm-
duct would be, if unwrapped from its coiling, of considerable
length; I found it to be gorged with sperm. The cirrus itself
was generally protruding from the genital aperture. It is not
large and has the usual structure of this organ. The difference
which it shows from the sperm-duct is very striking, and
furnishes an argument in favous of those who would hold that
the cirrus is not merely the end of the sperm-duct which is
capable of protrusion. It seems in this species at any rate to be
a structure independent of the sperm-duct. There is a certain
resemblance between the bottle-shaped cirrus sac and the bottle-
shaped receptaculum seminis and the end of the vagina. Both
of these tubes lie to the same side of the iateral excretory vessels —

68*

1000 MR. F. E. BEDDARD ON

if we are to regard the ovary as ventral, they lie to the dorsal
side.

The vaginal pore lies behind the opening of the male duct
into the genital cloaca. The vagina is a perfectly straight, delicate-
walled tube as in so many tapeworms. Posteriorly and close to
the lateral water-vessel, the vagina expands into a somewhat pear-
shaped receptaculum seminis, from the wider, posterior end of
which the vagina emerges again abruptly and runs a curved oblique
course towards the ventral side of the body. This latter region
of the vagina is of the same calibre and appearance as the
terminal section which opens into the genital] cloaca. The
swollen receptaculum seminis (text-fig. 206, p. 998) has on the other
hand thick glandular walls. The ovary and the yolk-gland he
anteriorly in each proglottid and very nearly in the middle of
the proglottid, verging however to the pore side, the position
being therefore quite different from that which characterises
Thysanotenia gambiana. The ovary is not large, neither is the
yolk-gland.

Whether a uterus exists as a definite structure at any period in
the development of the sexual organs, I am not able to state with
certainty. But I am disposed to think that a distinct uterus
does not exist. At the most, it must have a very brief existence,
for I can find nothing intermediate between eggs scattered in the
parenchyma and in the paruterine organs. J found nothing like
what has been described in Zhysanosoma. In this genus there
are stated to be outpocketings of the uterus round each of which
is formed a paruterine organ. I found in the present species a
condensation of the parenchyma round eggs or groups of eggs,
precisely as I have described in Thysanotenia gambiana. These
latter became more marked, and were then to be described as paru-
terine organs. In fact, the state of affairs which characterises the
genus Thysanotenia is to be looked upon as a further stage in
the development of such a genus as Oochoristica with the inter-
mediate formation of a uterus dropped out, or at least rendered
of very little importance. It is also like Davainea

Text-figure 207 represents a transverse section through a fully
mature proglottid of this worm. It may be compared with a
corresponding illustration of the other species of the genus,
Thysanotenia gambiana*, when certain differences will be
apparent coupled naturally with fundamental points of agreement.
In both species, the greater part of the medullary region of the
segment is occupied by the numerous paruterine organs, which
are only one layer deep. They extend between the excretory
vessels and up to those vessels on either side. In Thysanotenia
lemuris, however, the transverse row of paruterine sacs is at most
thirteen to sixteen, while they are much more numerous in
Th. gambiana. Furthermore, ina given section the greater number
by far—very often all—of the paruterine organs are seen to be

* P. ZS. 1911, text-fig. 158, p. 658.

NEW MAMMALIAN TAPEWORMS. 1001

- without a contained embryo or embryos. This does not mean,
of course, that these are paruterine organs which contain no
eges. It is simply an expression of the fact that in the present

Text-fig. 207.

Thysanotenia lemuris, transverse section through ripe proglottid, showing the
longitudinal external furrowing of the proglottid. e, paruterine organs, in only
two of which were eggs to be seen in this particular section; ¢, excretory
tubes.

1002 MR. F. E. BEDDARD ON

species each paruterus contains fewer eggs (or embryos) than
is the case with Thysanotenia gambiana. In no case have I
seen in a given section more than three embryos within
a single paruterine sac in the species which forms the
subject of the present communication. In reference to this par-
ticular I may compare text-figure 207 with the text-figure of
my paper dealing with ‘“ Thysanosoma ” gambianwm*. The
paruterine organs themselves were of about the same size in the
two species. The presence of so few embryos in a single par-
uterine sac produces naturally a very distinctive appearance which
at once distinguishes the two species from each other. In each
paruterine organ of 7h. lemuris it was possible to distinguish
a cortical and a medullary region of different appearance. ‘That
this was not possible in the other species is probably to be
accounted for by the large number of embryos which filled them.

It will, I think, be admitted that this tapeworm presents char-
acters which will not fit in with those of any known genus. It
contradicts indeed the definitions of families as given by Ransom,
at any rate to some extent; for I should be disposed to place
the genus in the neighbourhood of TZhysanosoma in the
family Anoplocephalide ; and yet this family is characterised by
the absence of a neck, present in the genus which is now under
consideration. The only other position in the series which this
worm could occupy, as I think, is in the subfamily Paruterine of
the family Hymenolepidide ; but in this subfamily the paruterine
organs are limited to one or two, and there is nothing like the
numerous organs met with in both of the species which I describe
here under the generic name of Vhysanotenia. This latter
reason as well as the unilateral genital pores prevent the inclusion
of the species in the genus Stilesia.

This new genus, Thysanotenia tT, may be thus defined :—

Thysanotenia, gen. nov.

Moderately large tapeworms, four to six inches in length and
three millimetres in breadth. Scolex unarmed, with laterally placed
suckers. Proglottids posteriorly as long as or slightly longer than broad.
Genital pores unilateral, borne upon a projection of the body near to
the anterior end of proglottids. Hxcretory tubes wide, one or two (and
these lying side by side) on each side, with or without ramified
branches. Reproductive organs a single set in each segment. Testes
numerous, or very numerous. Uterus a narrow transverse sac or
absent. Many paruterine organs in ripe segments. LEygs without
pyriform apparatus. Adults in Mammals.

* Loc. cit. text-fig. 159, p. 659.

+ Inasmuch as there are no fringes to the proglottids posteriorly such as occur in
Thysanosoma, the name selected is rather a misnomer. I have, however, used it
pore for the purpose of fixing what I consider to be the systematic position of
the worm,

NEW MAMMALIAN TAPEWORMS. 1003

Accepting the above as the generic characters of the genus
Thysanotenia, the two species may be thus defined :—

(1) Thysanotenia gambiana F. E. B.

Thysanosoma gambianum Beddard, P. Z. 8. 1911, p. 651.

Length about six inches, greatest diameter 6 millimetres. Segments
never longer than broad and only a few at the posterior end as lon;
as broad. Genital papilla not very conspicuous. Two excretory tubes
on each side, the inner of the two the larger, placed laterally
to each other ; a network of fine tubules connected with these. Testes
in two groups, the larger lying on the side furthest from the
genital pore, occupying the space between the two excretory tubes of
that side and a little beyond on each side. Ovary and yolk-gland on
pore side lying between the two excretory tubes and «a little to the
immer side also. Sperm-duct narrow or coiled, with a small vesicula
seminalis. No receptaculum seminis ; vagina opens into a terminal
muscular sac. Puruterine organs cach with many embryos. Uterus
a transverse sac.

Host, Gambian Pouched Rat (Cricetomys gambianus).

(2) Thysanotenia lemuris, sp. n.

Length four to sia inches with diameter of three millimetres. Seg-
ments at end of body rather longer than broad. Genital papilla very
conspicuous. One ewcretory tube on each side of body corresponding
to the innermost of the two present in Th. gambiana. No network of
tubules connected with this. Testes scatter ed through posterior part of
the body and anteriorly to the sides of ovary. Ovary and yolk-gland
submedian in position, slightly to pore side of segment. Sperm-duct
wide and coiled after issuing from cirrus sac. Lreceptaculum senrinis
present. Paruterine organs with only three or four embryos in each.
A uterus not formed (2). —

Host, Black Lemur (Lemur macaco).

It is obvious from the above definition and from what has been
said in the course of this paper that the two species, which I assign
to this new genus Thysanotenia, differ from each other in a
good many points, and perhaps may be considered to merit generic
separation.

Anoplotenia dasyuri, gen. et sp. n.

Atabout the same time, 7. e. from February 9th—11th, 1911, three
examples of the Tasmanian Devil (Dasyurus ursinus) died in the
Gardens ; only one of them was found to contain tapeworms, and
these occurred in that example in very great numbers. The
specimens belong, as I believe, to a new species, and I am also
disposed to form for it a new genus, which is to some extent allied
to the Anoplocephalide, but also, in the form of the uterus,
suggests Zwinia, sensu stricto. These facts have suggested the

1004 MR. F. E. BEDDARD ON

generic name which I here propose. I considered after a rough
examination of the worm that it might possibly be referred to
the genus Bertiella, and suggested this in a report to the
Prosectorial Committee; I am now convinced that it cannot be
included in that genus and that its characters will not allow of
its inclusion in any known genus. The movements of the living
worm were particularly active. The worm is not a long form and
has a rather unusually large head, as will be gathered from the

Text-fig. 208.

Anoplotenia dasyuri, enlarged about five times.

accompanying figure (text-fig. 208). I could find no trace of any
hooks or of any terminal sucker or other structure in the
rostellar region. The four suckers are unarmed and of large
size.

The scolex is well marked off from the strobila, although there
is no obvious neck—that is to say, the segmentation begins
apparently at onee. This is brought about not only by the

NEW MAMMALIAN TAPEWORMS, 1005

actually large size of the head but by the fact that the first strobila
are much narrower than those immediately following, and thus
an apparent constriction occurs which emphasizes the distinction
of the head. In other cases the conditions are apparently the
same; but an examination with a lens shows that though a
constriction at about the same distance from the rostellum exists,
the formation of strobila exists beyond this point and has invaded
the hinder region of the actual scolex. It is a little difficult
therefore to assert that a neck is or is not characteristic of
Anoplotenia dasyuri. The occasional commencement of
strobilisation immediately behind the scolex is not unsuggestive
of Oochoristica*, with which genus the present has some likeness,
and there is in the same way a kind of hint of a commencing
pseudoscolex.

The body of the worm is about an inch in length or sometimes
rather longer; but it never grows to a great size. The segments
which immediately follow the head are very short; but this
region of the body is not long, perhaps a couple of millimetres, and
contains but few segments, in some cases not more than a dozen,
in others rather more. The segments increase gradually in
length and towards the end of the body come to be three or four
times as long as they are broad. In the hinder segments the
genital pores are sometimes quite obvious when the worm is
examined with a lens, and irregularly alternate in position from
one side of the body to the other. In transverse sections the
body is apt to be hourglass-shaped owing to the greater thickness
of the lateral edges of the body due to the bulging caused by
the unusually large cirrus sacs. ‘This is certainly the case with
the shorter and flatter anterior proglottids.

In transverse sections, through the anterior region of the body
in mature segments, which are, however, not long and distended
with ova, the layers of the cortex can be readily distinguished.
The cortex (text-fig. 209) is of about the same diameter as the
medullary portion. It is distinguishable into a much thicker
outer layer of longitudinal fibres, and a much thinner inner
layer of longitudinal fibres. The two are separated by delicate
transversely-running fibres, of which there are also a set
within the inner layer of longitudinal fibres and thus bordering
upon the medulla. The inner layer of longitudinal fibres is
particularly conspicuous for the reason that several are closely
grouped into a bundle, of which bundles there is only a single
row, as is shown in text-figure 209. This arrangement of the
muscular fibres is only apparent in the more anterior segments.
It ceases to be obvious in ripe proglottids such as that represented
in text-fig. 213 (p. 1012) where the body is gorged with eggs.
In these segments however, where, as will be pointed out at
length presently, the eggs are partly contained in a uterus and
partly scattered singly or in groups through the medullary

* See P. Z. S. 1911, p. 628.

1006 MR. F. E. BEDDARD ON

parenchyma, a new set of muscular fibres becomes apparent which
I have not observed in the anterior segments and which therefore
if really present are not so plain in those segments. The fibres run.
across the medullary parenchyma in a dorgo-ventral direction and
apparently belong to the cortical layer of circular fibres. In
text-fig. 213 some of these fibres are seen to cross the medullary
parenchyma and then to join the longitudinal series of cortical
fibres. Possibly these fibres assist in the rupture of the
proglottids to expel the eggs, and their presence may also account
for the particularly active movements of this species which have
been referred to above.

Text-fig. 209.

Anoplotenia dasyur?, transverse section through part of body-wall. I, two delicate
layers of circular fibres between which is a special layer of longitudinal fibres
grouped into bundles. 7, testis.

The excretory vessels are two on each side and are accurately
superposed, 7. e. dorsal and ventral. Later on in the body only one
vessel is very plainly visible on each side, and this is of greater
calibre than anteriorly.

The gonads and their ducts appear very early in the strobila of
Anoplotenia dasyuri. They commence to be visible within less
than 1 mm. of the head immediately after the anterior set of very
short segments. ‘The segments, when the ducts are first visible,

NEW MAMMALIAN TAPEWORMS, 1007

are absolutely crammed with apparently mature (certainly very
nearly mature) testes. The ducts showed no signs of specialisation.
The vas deferens only expanded slightly and gradually into the
elongated terminal sac, which is so much specialised in the mature
segments to be described later, This immature condition of the
ducts persisted for only six segments. Thereafter the cirrus sacs
were nearly or quite fully developed. In these anterior segments
the ovaries were not so forward in development as the testes.

Text-fig. 210.

Anoplotenia dasyuri, longitudinal section of proglottid. A., shell-gland; Ov., ovary,
below which is seen the cirrus sac ; 7’., testes ; U., uterus, between which and the
cirrus sac are seen the coils of the vas deferens cut transversely ; W., transverse
excretory tube. The posterior part of the proglottid is above.

The ovaries of Anoplotenia dasyuri lie posteriorly in the
segment but anteriorly to the vitelline glands, and when fully
developed are large and distinctly double. They have the very
common bushy form and occupy a good deal of the posterior
region of the segment. I did not observe the ovaries to be fully
developed until the first segment, in which the cirrus sac is also
fully developed and in which the uterus has begun to appear.
When the uterus has become so far developed as to fill the greater
part of the proglottid, the ovaries quite distinctly lie in continuity
with the masses of not fully mature eggs which here fill up the
chambers of the uterus.

The vitelline glands are quite evident close to the posterior
border of the proglottid; they do not extend out so far laterally
as do the ovaries.

The vagina of this worm is, as is so usual in the group, a quite
straight tube for the greater part of its extent, that is to say it
is not coiled. It lies behind the cirrus sac and opens into the

1008 MR. F, E. BEDDARD ON

commencement of the genital cloaca, perforating the muscular
pad. Its walls are not distinctly cellular and they stain deepiy,
both of which features are very common in these animals.
Distally, the vagina may be easily followed until it opens into a
well-marked receptaculum seminis, which is very large and con-
spicuous in Anoplotenia. In young segments its course is straight
across the segment; in older ones it passes straight to the
middle line in an oblique course, then bends back along the middle
line of the segment. This sac lies almost exactly in the middle
of the body and is absolutely circular in transverse sections of the
proglottids. ‘The vagina enters it on the ventral surface and
leaves it again at an exactly corresponding point further towards
the non-pore side of the proglottid. Thus the tube leaving the
receptaculum has to be followed for a short distance in sections
before it can be ascertained whether it is the distal or proximal
part of the vagina. The receptaculum was gorged with sperm. It
is not spherical as might be imagined from its circular contour in
section, but narrows to the more slender tube at one end which is
beyond the exit of the vagina.

Towards its point of opening into the genital cloaca, the vagina
lies parallel to the cirrus sac, in some cases being less oblique
in its course than in other proglottids. It has a very thick
muscular wall, that is to say thick relatively to the size of its very
small lumen. This muscular coat is double, an inner longitudinal
and an outer circular layer being present. Opposite the actual
point of opening into the genital cloaca, the vagina suddenly
widens into a very small sac lying closely adpressed to the
muscular pad which forms part of the wall of the genital cloaca ;
from this sac, a narrow tube passes at right angles to the
rest of the vagina and perforates the muscular pad. The terminal
sac of the vagina is seen to be filled with sperm. It is related
perhaps to the lateral orifice of the cirrus within the invaginated
pouch of the cirrus sac: this when protruded forms a bulbous
extremity, near to which the orifice would pour its contents
into, quite fill and perhaps even somewhat dilate, this terminal
sac of the vagina. This matter is, however, more fully dealt with
under my description of the cirrus sac and penis (on p. 1014).

I am disposed to think that the receptaculum seminis is no
more than a dilatation upon the vagina, for it could easily owe its
shape to mere gorging with sperm, and its walls appear to be like
those of the rest of the vagina and to have lost their cellular
character. It is at any rate greatly disguised in them as in so
many other tapeworms. As Gough has lately pointed out,* this
non-cellular appearance is preceded by a distinct wall of cells. If
the receptaculum seminis be as I suggest merely a local swelling
of the vagina, it is clearly quite different in its nature from the
receptaculum described above in Zhysanotenia lemuris. ‘This
latter is most obviously a distinct and definitely specialised region

* “Tapeworms of the subfamily Avitelline,”’ Quart. Journ. Micr: Sei. vol. lvi.

NEW MAMMALIAN TAPEWORMS. 1009

of the female tubes. For it is apparent in less mature proglottids
and has there walls of a different character from the slender

Text-figc. 211.

Anoplotenia dasyuri, two vipe proglottids viewed as transparent objects.
L., masses of eggs at posterior end of proglottid; R., reticular portion of uterus ;
Sp.d., cirrus sac.

1010 MR. F. E. BEDDARD ON

vagina connected with it. And, moreover, in these younger
proglottids its shape cannot be due to any distention by sperm,
for the sac was quite empty of sperm. It might be con-
venient to restrict the term receptaculum seminis for cases
of this kind, and not perhaps to name specially the distended
region of the vagina in which the sperm is chiefly massed in
other forms.

Text-fig. 212.

Anoplotenia dasyuri, transverse section through ripe proglottid.
O.,ova; U., cavity of uterus.

The uterus of this tapeworm is visible very early in the body
in correlation with the early development of the organs of repro-

NEW MAMMALIAN TAPEWORMS. 1011

duction generally. I found, in fact, that the uterus was quite
recognisable in the first segment which had a fully developed
cirrus pouch, and that segment was one of the earliest to have
attained an appreciable length and lay hardly a millimetre
behind the scolex. In this segment the uterus showed (in a
longitudinal horizontal section through the anterior region of the
body) a rounded form stretched in the direction of the transverse
section of the body and thus rather oval in outline, and it
occupied precisely the median region of the proglottid. In this
particular uterus I found no ova. A segment or two further
back the uterus is already larger, but it still has the form of a
more or less oval sac, extending in these segments towards the
pore-side and having thus become eccentric in position. The
eccentricity, however, is not very strongly marked. In these
segments the uteri were full of ova.

"The uterus in these segments lay near to the posterior boundary
of the proglottid and was transversely elongated in form; it
was distinctly posterior to the strong muscular cirrus sac. The
uterus possessed a distinct epithelial wall that was of sufficient
thickness to show itself in all my sections (cf. text-fig. 210, p. 1007).
Later, the epithelial wall is not obvious, but the cavity has plain
boundaries and can be recognised as a definite cavity and not
merely a system of irregular lacune.

In the posterior ripe proglottids the uterus undergoes some
changes which are not altogether easy to follow and to correlate.
When the elongated, fully ripe proglotiids are examined mounted
in glycerine, the eggs are seen to be arranged throughout them
in a way which differs slightly in different proglottids but
is as arule at any rate on the same plan. ‘The eggs occur in
clusters and strings which give the appearance of a retiform
uterus. In this, at times, a median string of eggs giving off
lateral branches may be recognised. And though these lateral
branches join here and there and thus make a network, the
general appearance given in such segments is that of the uterus
of 7 cenia, Which is characterised ‘rs a median stem and lateral
branches. Very commonly the ova are more thickly clustered
together in the posterior region of each segment. There is, I
think, little doubt that if the worm were examined only in this
way, the uterus would be pronounced to be reticular. A study
of sections, however, leads to a rather different interpretation of
the arrangements visible in solid preparations. In some trans-
verse sections, such as that illustrated in text-fig. 212, the whole of
the interior of the proglottid is occupied by the uterus and the
contained masses of developing eggs. These appear to lie in a
large undivided cavity, which I take to be the uterus. This
region corresponds to the posterior part of the segment, where as
already mentioned the eggs tend to become massed.

In other sections through the same proglottid as that which
has just been referred to, the conditions observable were different.
There are, as is shown in the accompanying figure, eggs and

1012 MR. F. E. BEDDARD ON

groups of eggs which correspond to the thinner strings of eggs
in proglottids which are viewed as solid objects rendered trans-
parent by glycerine. These groups vary in size, but it would
appear that they are imbedded in the parenchyma of the medul-
lary region and are not contained in cavities—that, in fact, there

Text-fig. 213.

Anoplotenia dasyuri, transverse section through ripe proglottid.
O., ova, contained in uterus (U.); O1, eggs scattered through parenchyma.

is no uterus here at all. It may, of course, have been present
and have disappeared. In intermediate proglottids the uterus
forms a series of cavities which apparently intercommunicate and
thus constitute a network. In these cavities the eggs are not
mature—at any rate, the membranes have not yet appeared.
This system of cavities fills up a great deal of the available space

NEW MAMMALIAN TAPEWORMS. 1013

and the testes become much restricted and tend to disappear.
The most noticeable part of the medullary tissue left is a pro-
jection from the region of the cirrus sac lodging the coil of the
vas deferens.

It would appear therefore that the uterus in this genus
Anoplotenia rather combines the characters of that organ in
several other genera than presents us with a new type. That it
passes through a simple saccular stage is not perhaps a fact of
any great moment; for that occurs in many genera. But it 1s
undoubtedly reticular at one time, and, contrary to what is found
in such cases, the reticular stage is not permanent.

Text-fig. 214.
RS.

Anoplotenia dasyuri, longitudinal section through proglottid, showing the branched
and reticular uterus with ovain smaller and larger clumps. B.S., receptaculum
seminis; W., excretory tube.

The ultimate condition of the uterus does not fall definitely
within any of the types used by Ransom in his table of arrange-
ment of the genera of Teenioid Cestodes ; and the condition of this
organ in Anoplotenia dasyuri is really one of the chief reasons
upon which I base its generic distinction from other forms.

The ¢estes of this species are very numerous in those anterior
segments in which they are at their full development. They
are pressed closely together and overlap and appear to fill all of
the available space left between the ovaries and other organs of the
proglottid. As the latter are posterior in the segment, the testes
are mainly anterior. Both in longitudinal and sagittal segments
the testes can be seen to be limited in their occurience only by

Proc. Zoot, Soc.—1911, No. LXIX., eo

1014 MR. F. E. BEDDARD ON

the cortex. In correlation with their large numbers, the testes
are of small size. They cannot, obviously, be said to be either
dorsal or ventral or anterior or posterior in position.

A good deal of the anterior part of each mature proglottid is
occupied by a large coil of the vas deferens which forms a larger
mass than in many tapeworms figured or known to me at first
hand. Although this coil lies anteriorly in the segment on a
level with the lar ge cirrus sac, it does not touch the anterior
boundary of the segment. In front of it there is to be seen a
considerable heap of testes. The coil is generally in close contact
with the cirrus sac. I found no vesicula seminalis in this species,
but the coiled tube gets wider when ripe.

The cirrus sac and its contained structures are rather re-
markable in this species and much more complicated than in
many other species, including the three that I have already
described in my former communication *. As already stated,
the organs of reproduction appear very early in the chain of
proglottids. It is, however, not for some segments that the
cirrus sac is fully developed. The fully developed cirrus sac
coincides with the first appearance of the uterus. As already
mentioned, in considering the external characters of this worm,
the cirrus sac when mature is so large as to cause an appreciable
bulge in the segment. It therefore entirely fills the medullary
region of that part of the segment where it occurs, as seen in
a transverse section. In such sections it may also be seen
that the cirrus sac and the genital cloaca together (of which
a description follows) occupy about one-third of the entire breadth
of a proglottid.

The cirrus sac is nearly if not quite spherical in shape, and
consists of an outer coat and of an inner mass of tissue. The
outer coat is not very thick and is muscular, the fibres no doubt
serving to compress the sac and thus bring about the evagi-
nation of the cirrus. The internal tissue of the cirrus sac is
a tissue in which the cirrus itself is embedded, and it entirely
fills the sac save where it is traversed by the cirrus. It contains
many nuclei scattered fairly closely throughout it, and delicate
fibres which may be occasionally seen to possess a distinctly
retiform arrangement. It seems to me to be a soft tissue which
is an elastic packing material transmitting to the cirrus the
contractions of the external muscular coat of the cirrus sac. In
some sections indeed the nuclei can be seen to be more compressed
and regularly arranged in lines in the immediate neighbourhood
of the eversible cirrus sac. This might seem to argue some
contractility possessed by the tissue forming the core of the
cirrus sac.

The cirrus itself is peculiar and complicated in structure. It
consists of two parts. First of all there is the part which lies
immediately within the cirrus sac and which is perfectly con-
tinuous with the vas deferens, and shows no sudden differences

* P.Z. S. 1911, p. 626.

NEW MAMMALIAN TAPEWORMS. 1015

of structure that I can detect from the vas deferens. It lies in
a loose coil of only two or threeturns. Followed distally, this tube

Text-fig. 215.

Anoplotenia dasyuri. Cirrus sac with penis in various stages of retraction.

In the upper figure the penis is completely retracted, in the middle figure it:
is completely protruded; in the lower figure it is incompletely protruded.
A, muscular pad on anterior side of genital cloaca; B, muscular pad on posterior
side of genital cloaca which is perforated by opening of vagina (Ja.) ; C., cirrus;
P., penis, on the posterior side ot which, as is shown in the middle figure, the
cirrus opens; Sp.d., vas deferens.
(SiS)

1016 MR. F. E. BEDDARD ON

is seen to open into an invaginated sae which lies pushed into the
‘solid core of the cirrus sac, as is shown in the text-figure accom-
panying this description. This sac is irregular in form with
crumpled walls, and it is surrounded by a layer of particularly
stout muscular fibres, which lie therefore within the cirrus sac
and form a differentiated portion of its core. These muscular
fibres lie loosely round the invaginated sac. The wall of the
latter is rather thickened to form a pad lying on the anterior
side, and close to this the cirrus opens into it, their cavities
becoming continuous. The opening into the sac is opposite to
the pad, and therefore on the posterior side of the sac.

The cirrus sac does not open directly on to the exterior, but
through a genital cloaca which is itself much complicated. <A
horizontal section through the whole structure is represented in
text-fig. 215. The genital cloaca may be divided into three or
four regions; the invaginated penis (as we may term the in-
and evaginable sac into which the cirrus opens) is continuous
with a wide but narrow cavity of quite as great a diameter
(antero-posterior) as the cirrus sac itself, but very narrow from
side to side. After this comes a tubular cavity surrounded
by a very thick layer of circular fibres which are perforated, as
already mentioned, on the posterior side by the vagina which
here enters the genital cloaca. In horizontal sections such as that
represented in text-fig. 215, this layer of muscles appears as two
strong muscular pads, that on the posterior side being longer
from side to side than the one opposite. After this the canal
widens a little for the terminal part of its course and its walls
are rather crumpled, there being a particularly deep recess”
immediately after the sphincter region which precedes it. A
considerable portion of the cirrus sac can be evaginated, sometimes
more and sometimes less. These differences also are shown in
text-figure 215. In some cases only the anterior side of the
cirrus sac is protruded, which in such a case barely reaches
the external orifice of the genital canal. In other cases
much more is protruded, and the cirrus itself is drawn down
into this penial protrusion which extends well beyond the
external orifice. In this latter case, however, which represents
the extreme of what I have seen in my sections, the actual opening
of the vas deferens is not at the tip of the protruded penis, but
at the side and within the genital canal. I may not perhaps have
seen instances of extreme protrusion. It is noteworthy that the
orifice of the vas deferens is on the posterior side of the penis and
thus corresponds to the vaginal orifice. Possibly a complete
extrusion of the penis takes place in cases of cross-fertilisation.

The following are the general characters of this species * :—

Head quite unarmed, of rather large size and with four unarmed
suckers, Strobilisation begins at once, there being no “neck”: the
jirst two or three strobila wider than those which immediately follow

* IT do not attempt to discriminate between generic and specific characters.

NEW MAMMALIAN TAPEWORMS. 1017

and of same diameter as or wider than head, thus forming «a
rudimentary pseudoscolex. The proylottids increase very rapidly to
a consideruble length, the posterior beiny longer than broad and
becoming detached. Not more than ten or a dozen anterior short
proylottids. Genital orifices single and irregularly alternate.
Exeretory tubes posteriorly one on each side of body, that of one side
being as a rule wider than that of opposite side; in anterior segments
two on each side. Testes very numerous, filling up the whole space
left by other organs in proylottid. Vas deferens coiled. Cirrus sac very
large and spherical and somewhat peculiar in structure, with an
eversible sac reaching the exterior through a much differentiated
genital cloaca. Ovarres posterior in segment and with vitelline glands
posterior to these. Vagina straiyht and narrow, opening posteriorly
to currus sac; a receptaculum semuus present. Uterus at first a
simple sac, later a reticulum, and later still part of the uterus remains,
while other eggs are imbedded singly or in groups in the medullary
parenchyma. Hggs without V-shaped apparatus.

It will be, as I think, evident from the résume of the characters
of this species just given, that it cannot be referred with any
confidence to any one of the really known genera of the Tetra-
cotylea. The convenient table giving a key to the various genera
used by Ransom in his memoir enables one to refer the species
from Dasyurus ursinus to the neighbourhood of Oochoristica, Tenia
(s.s.) or Bertiella. Of the latter genus several species are known
from Marsupials; but they are not known from the present
genus, and appear to be nearly limited to the herbivorous (at
any rate Diprotodont) genera, i.e. Phalanger, Phalangista, and
Phascolarctos*.

These species, however, are certainly not congeneric with that
which I describe in the present paper. They agree with the
generic definition given by Ransom7, who doubtless took them, as
well as the species of Bertiella from Apes and Rodents and Birds,
into consideration when formulating his definition. The worms
studied by myself show the following important differences from
Bertiella as defined by Ransom :—The strobilisation is different,
the posterior strobila being much longer than broad ; the genital
canals pass between the dorsal and ventral excretory vessels ;
the testes exist throughout the segment save where space is
occupied by the ovaries etc.; the uterus is of a totally different
character; the cirrus sac is also totally different from anything
figured in Bertiella. I do not feel able therefore to refer this
species from Dasyurus ursinus to the genus Bertiella.

I am of opinion that the present genus is nearer to the genus
Oochoristica. The latter genus actually occurs in carnivorous
Marsupials but in Neotropical forms, in fact in Didelphys, and not,
however, so far as I am aware, in Australian Marsupials. The

* See Zschokke in Semon’s ‘ Reise,’ Jena 1898, for B. obesa and B. semoni; and
the same author, “ Neue Studien an Cestoden aplacentaler Siugethiere,” Zeitschr.
wiss. Zool. lxv. 1899, for B. edulis and B. sarasinorum. Also Janicki, “ Die
Cestoden Neu Guinea’s ” in Nova Guinea, Livy. v., 1906, p. 281, for B. rigida.

+ Loe. cit. p. 62.

1018 MISS R. HARRISON AND PROF. S. J. HICKSON ON

general form of the body and the segmentation is not unlike in
the two genera; and especially to be noted is a resemblance in
the scolex. In the present genus as in the Oochoristica from
Tamandua tetradactyla, described by myself * some months since,
the strobila, as it were, invade the scolex. On the other hand, the
early disappearance of the uterus and the imbedding of the ova.
singly in the medullary parenchyma is a character of Oochoristica
which distinguishes it from the genus which I propose to call
Anoplotenia. The peculiar cirrus sac and the very complex
genital cloaca are points in which Anoplotenia differs from all the
genera with which I here compare it.

There now remains the genus Tenia (sensu stricto) to which the
present species shows a certain amount of likeness in the uterus,
which is rather pronounced in certain proglottids. There is in
fact occasionally a quite distinct median stem with branches.
Tenia, however, has an armed rostellum which is sometimes not
armed asin 7’. (eniarhynchus) saginata, where the hooks drop out
early and are replaced by a sucker-like structurey. There is
nothing of this kind in the present species, which moreover bears
no such close likeness to Zenia saginata as would warrant its
inclusion in the same genus or subgenus. Another genus in
which the uterus has a marked median stem and lateral branches
is Catenotenia,: the species of which occur in the mouse and in
the squirrel. in this genus, however, the testes and ovaries have
a different position from that which is met with in the tapeworm
dealt with in the present memoir, and the relation of the genital
duct to the excretory tubes is also different.

46. Some Madreporaria from the Persian Gulf. By Ruts
Harrison, Oxford §. With a Note on the Memoir and
some Further Notes on Pyrophyllia inflata by SyDNEY
J. Hickson, M.A., D.Sc., F.R.S., F.Z.S.

[Received May 19, 1911: Read June 27, 1911. ]
(Plates LVIT. & LVIITI. |! and Text-figures 216-221).

This collection of Madreporarian corais was made by Mr. F. W.
Townsend, and entrusted to me for identification and description
by Professor Hickson. I should like to take this opportunity of
thanking Professor Hickson for putting this interesting piece of
work in my hands. My thanks are also due to Professor Bourne
for allowing me to carry on the work in his laboratory and placing
all its resources at my disposal, and for help and advice during the

* P. Z.S. 1911, p. 627. I ought to have mentioned in that paper that something
of the same kind appvears to occur in O. rostellata (see Zschokke, Zeitschr. wiss.
Zool. vol. Ixxxini. 1905).

i Cf. Bronn’s “'Thierreich,” Vol. iv. Abth. B. p. 1720.

{£ Janicki, Zeitschr. wiss. Zool. 1906, vol. Ixxxi. p. 505.

§ Communicated by Prof. S. J. Hickson, E.R.S., F.Z.S.

|| For explanation of the Plates see p. 1044.

1% Bots) MSIL, PW IOWA,

MADREPORARIA FROM THE PERSIAN GULF.

I, & S, Ill, il. ISVAN,

pis Paina ee roy

poster

MADREPORARIA FROM THE PERSIAN GULF.

Bale & Danielsson Lttim;

i,

%

CORALS FROM THE PERSIAN GULF. 1019

progress of the work; to Professor Sollas for permission to use
his apparatus, which enabled me to grind sections, photograph and
reconstruct a wax model of Zrematotrochus zelandic ; to Professor
Jeffrey Bell for permission to examine various Madreporaria in
the British Museum; and to Professor Stanley Gardiner for the
loan of a large number of his specimens of Heterocyathus
. equicostatus for comparison with the present collection.

The collection consists of examples of the following species :—

Family GuyNnrp&.
Pyrophyllia inflata Hickson [25].
Family FLABELLIDS.
Flabellum magnificum v. Marenzeller [28].

Family TurBInoLup.

Heterocyathus cequicostatus Milne-EKdwards &
Haime [29].

Heterocyathus heterocostatus, sp. n.

Paracyathus cavatus Alcock [1}.

Trematotrochus zelandie Dunean {14}.

Agelecyathus persicus Duncan |14 |.

Family Funerp.
Fungia patella Milne-Edwards & Haime [29].

Family HupsaAMMUID&.

Heteropsammia aphrodes Alcock |1}.
Dendrophyllia sp. ¢

All these specimens are in the Manchester Museum. The
occurrence of a recent Zrematotrochus is worthy of special
attention. Hitherto, the genus has been known from seven
fossil and one recent species from Australia and Australasian seas,
and it is remarkable to find it appearing in so remote a locality
as the Persian Gulf. More remarkable still is the fact that this
very species has already been described from Cook’s Strait,
New Zealand, by Professor Martin Duncan under the name of
Conocyathus zelandie. 'The resemblance of the coral I have been
examining to the figures and description (so far as it went) of
this Conocyathus was so striking, that it occurred to me that
possibly Duncan had overlooked the perforations of the wall,
characteristic of a Trematotrochus. Owing to the kindness of
Professor Jeffrey Bell I have been able to re-examine the original
type specimens in the British Museum, and the result of this
examination has been to convince me that the corals are absolutely
identical. ‘The British Museum species have not been so care-
fully cleaned and dried as those in the present collection, and in
places the intercostal furrows are somewhat choked up by sand
and grit; but that perforations do exist, exactly similar to those
of the Persian Gulf species, | have no hesitation in stating ; in
parts of the corallum they are clearly visible, but unless one were

1020 MISS R. HARRISON AND PROF. S. J. HICKSON ON

expecting to find them it would be very easy to overlook them,
and it is not altogether surprising that Duncan should have done
so. Had he observed them, I venture to think he would not have
called the coral a Conocyathus. In his ‘ Revision of Families and
Genera’ he places Conocyathus, Trematotrochus, and Turbinolia
together as closely allied forms; indeed the perforations of the
theca of Trematotrochus form the only feature which separates it
from Conocyathus. Now that perforations have been observed in
Conocyathus zelandiw, we must either amend the definition of that
genus, or else remove this species to the genus Zvrematotrochus.
The existence of pores is a character of such importance that the
latter course seems to me advisable, and in future Conocyathus
zelandie should be known as Z'rematotrochus zelandiwe. The
specific name is unfortunate and apt to be misleading, as is
bound to be the case when a specimen is given a name denoting
the locality in which it was originally found, and subsequently
appears in other parts of the world. A further consideration of
the structure and systematic position of this coral will be found
in the systematic part of this paper.

Family GUYNIID&.

PyROPHYLLIA INFLATA Hickson [25]. (Pl. LVII. figs. 8-11;
Pl. LVIII. figs. 18, 19.)

About sixty specimens of this species were obtained on a
gravelly bottom at a depth of 156 fathoms in the Gulf of Oman.
Professor Hickson has added some further notes to his original
description of this species at the end of this memoir (p. 1039).

FLABELLUM MAGNIFICUM v. Marenzeller [28.] (Pl. LVII.
figs. 1-3.)

Corallum fan-shaped, wall and septa very thin and delicate.
Numerous rootlets descend vertically downwards, each such
rootlet communicating with two interseptal chambers on opposite
sides of a septum, and firmly fixed on a mass of mud and
serpulid tubes. Calice oval in outline, but constricted in the
middle of the short diameter; practically semi-circular at the
ends of the long axis. ‘Two specimens, measuring as follows :—

I. II.
1a (ytd Ni Ave peeer eee ral, eh aan se aenre 2Suailicee Hy ees SELON TT Ta 15 mm.
N Rio ar 6) LOOM CVENTACIES socooausocnvooonndve seconn eon 60 . 40
Calice ry Shoitidiamerers ‘ 4
‘ Greatest width ..............000000. 40 ,, 26 ,,
Least width? ees fees. aoO) see Zowes
Numibertofiseptaywen.ea ee eer cee eee rela 96 |
As 6p DUNT SEI, saocodondenocoouocedenaact 24, 24 |

CORALS FROM THE PERSIAN GULF. 1021

Costee faintly indicated throughout entire height of the corallum
which is also marked with fine wavy transverse accretion lines.
Septa in six systems of six cycles of which the sixth cycle is
incomplete, while there are present a few rudimentary septa of
a seventh cycle. Those of the first three cycles equal and reaching
the columella, edges vertical, entire until within about 4 mm. of
the columella where a few coarse denticulations may be present ;
beset with minute blunt spines arranged in transverse and radial
rows. Septa of lower cycles become successively smaller. Columella
parietal, only slightly developed.

Locality. Telegraph Cable, Persian Gulf. Depth not recorded.

In the larger specimen there are 175 septa arranged as
follows *:—

Bhi " | Number of super-
Number of 7 unter of sels of numerary septa of
septa. 1 cycle wanting. 7th cycle.
Chambers] aier.cseereeet 31 | 3 2
Be 2) tec eae 25 7 none
a) Wubeene ecdane 30 2 none
|
EL. 26 | 6 none
a 5 30 | 2 none
Gite shove ike 33 5 6
otaliaesee scenes 175 25 8

This coral differs from v. Marenzeller’s species in its smaller size
and the presence of numerous rootlets. Gardiner [19] has shown
that the presence or absence of such rootlets is a variable
character, and he has described rootlets in Flabellum rubrum, a
species in which they had not previously been recognized.

v. Marenzeller’s species was made for a single specimen, and as
the agreement between the calicular and septal characters of the
specimens under consideration and those of the type are so
similar, it has been considered advisable not to separate them.

Family TURBINOLIID4.

Genus HETEROCYATHUS.

The genus Heterocyathus has been critically examined by
Gardiner [20], who had at his disposal a very large number
of specimens. He absorbed the species of Semper and Rehberg
in a single variable species originally described by Milne-

* The septa are considered as being divided into six chambers, bounded by

primary septa; the chambers are considered in rotation beginning with one to the
right-hand side of a directive septum.

1022 MISS R. HARRISON AND PROF. S. J. HICKSON ON

Edwards & Haime [22] as Heterocyathus cwquicostatus ; to this
Bourne [6] has since added Stephanoseris rowsseawi. .

In the large collection of corals from 8. Africa, Gardiner
recognized two types with the following diagnostic characters :—

Type I. Coste equal in size, rounded and covered with low
granules (Pl. LVIILI. fig. 12), small intercostal spaces ; base smooth,
low granules, only traces of cost ; theca with thin upper edge,
never more than 1 mm. above columella; septa in four cycles,
thick with narrow interseptal spaces, average exsertness 1°5 mm. ;
cycle i. broader and more exsert than cycle 11., 11. than ii1., iv.
more exsert than ii1., higher on either side of i. than of 11. Sides set
with low ridges, edges not toothed ; commonly certain ones, or all,
coloured black; pali before all cycles, large and conspicuous ;
columella a mass of rods decreasing in size and height from
the pali towards the centre of the axial fossa, densely packed
together.

Tyre 11. Coste of cycles i., ii., and ii. larger than iv., which
consists of a row of separate granules rougher and higher than in
Type I, with broader intercostal spaces (Pl. LVIII. fig. 13); base
roughly granular, costee sometimes extending on to it; theca with
thin upper edge, generally 2 to 3 mm. above the top of the
columella; septa in four cycles with a tendency to have some of
a fifth cycle represented, thin with broad interseptal spaces,
average exsertness 3 mm.; ridges on sides conspicuous, edges
toothed towards centre of calice; all the same colour, white or
some shade of grey; no proper pali or columella; septa iv. fuse
with septa ili., and these again with 11. ; septa 1. generally separate,
but in the centre fuse with the rest forming a mass of trabecule
covered by fine points which run up along the septa of cycles 1. to
ui. for some distance, almost like fine teeth.

In describing the collection of Mr. J. J. Simpson and
Dr. Rudmose-Brown from Burma, Miss Poole [24] recognized
the Gardiner’s two types, and added a third type with a fifth
eycle of septa, four crowns of nodular pali, and a deeper fossa.

I have been able to re-examine the collections described
by Professor Gardiner and Miss Poole, and, so far as the former
collection is concerned, the two types are sufficiently different,
in my opinion, to be regarded as two species. Gardiner himself
has. described them as “ two perfectly distinct modes of growth,
almost two varieties.” I have searched in vain for intermediates,
and although I had no difficulty in picking out the two specimens
which the author describes as doubtful, they are both too much
broken and corroded to afford any clear evidence that they
vepresent a form intermediate between the two types. The
general facies are entirely different. In Type I, the tendency is
for the base to be broader than the calice in correlation with
rounded granular coste, thick septa, practically no fossa, and a
papilliform well-developed columella. In Type I, the tendency
is for the coral to taper away toa point from a circular calice ;
such is the form of the only known example without a commensal

CORALS FROM THE PERSIAN GULF. 1023

Aspidosiphon; the flat base of the normal specimen is never so
large as the calice, and this shape is invariably in correlation with
alternating coste, thin septa, a deep fossa, and an ill-defined
columella.

The characters of the coste of the two types are very definite.
Fig. 12 on Plate LVIII. represents two adjacent cost of Type I,
low broad ridges, covered all over with fine uniform granules; the
intercostal furrows are small and shallow. Fig. 13 on Plate LVIII.
represents two adjacent cost of Type Il: on the right is the
type of costa which corresponds to septal cycles i., 1i., and 111.,
a prominent exsert ridge, on which the granulations tend
to become regular transverse bars; on the left is a costa corres-
ponding to septal cycle iv., slightly less exsert, and beset with
very irregular coarse granulations; the intercostal furrows are
deep. These marked differences, always in correlation with the
differences of the calicular characters to which reference has
already been made, are too consistent to retain both forms in the
same species, True, there are many examples of Type I which
have an alternation of broad and narrow coste, but such coste are
always alike in being lowand uniformly granular; the alternation
of coarsely granular coste and much exsert narrow coste is never
discernible. In some, however, there is a tendency for the coste
of Type I to become more exsert and more coarsely granular at
the calicular margin, but this generally occurs where the whole
growth of the coral has been distorted by being fixed to an
abnormally large shell, and 1t might be thought that this distortion
had influenced the form of growth. Indeed, it suggested the
possibility that the shell on which the coral fixes itself, and the
position which that shell assumes within the actual body of the
coral, might influence the mode of growth to such an extent
that not only the general shape but the character of the septa
and coste might be controlled. This, however, is not the case.
There are well-defined examples of Types I and IL both fixed
on exactly similar shells: in some the Aspidosiphon in corals of
both types is coiled horizontally forming a flat base, in others the
corals are fixed to shells which le sometimes horizontally,
sometimes vertically within the coral zoophyte, and there are
always examples of both types harbouring Aspidosiphons in shells
which assume either position; so that the species of shell or
the position it assumes exercises no influence on the manner of
growth of these two well-defined types. Such types vary about
two distinct modes, with a slight overlapping of the extremes of
variation ; and are not themselves the extremes of a single growth
mode. The relation of the species to each other is represented in
text-fig. 216.

Of some 252 individuals in the South African collection, there
are 225 of Type I and 19 of Type IJ,and 8 which have characters
commonto both types. The characters which separate the two types

cannot be Bees and given numerical values ; consequently
in text-fig. 216 A the point about which the two growth-modes

1024 MISS R. HARRISON AND PROF. S. J. HICKSON ON

vary is purely arbitrary, but the number of individuals which vary
about these points is a real number, and represents the two
distinct growth-modes of two different species. If the two types
were varieties of a single growth-mode, the variation might be
expressed by such a diagram as text-fig. 216 B, but that would
represent a condition of affairs wholly contrary to the numerical
facts; for the intermediates, which in reality are less than a
thirtieth of the whole number, here represent the greater number
of forms.

Text-fig. 216.

() 20 40 60 80 100 420 140 160 180 200 220 240 260

ty) 20 40 60 go 100 120 140 /60 180 200 220 240 260

A.—A diagrammatic representation of two species which vary about two distinct modes,
the extremes of which converge towards one another. The abscisse give numbers
which vary about each mode respectively.

—A diagr: atic representation of a single species varyin 0 i inar ,
B.—A diagrammatic representation of a singl ar about an imaginary mode

Type I is undoubtedly the H. equicostatus of Milne-Edwards
& Haime, although, as I shall show, there are considerable
variations within the type. Type II is probably identical with
Verrill’s [42] H. alternatus, a species which has escaped the
notice of several authors. This species possesses all the
characters which separate Gardiner’s Type II from Type I,—the
base smaller than the disk, with a slight constriction above it, and
then walls spreading obliquely outwards to the edge of the disk,

CORALS FROM THE PERSIAN GULF, 1025

alternating costz, paliform teeth exsert before all septal cycles
except the last, and an ill-developed papillose columella which
scarcely rises above the surface of the broad shallow central fossa.

When Miss Poole stated that the forms in the Burmese
collection belonged to three different Types, two of which were
identical with Gardiner’s Types I and II,she had not the advantage
of seeing the actual specimens, and was in error. All the
Burmese forms belong to Gardiner’s type I, but they show great
variation, and in order to justify their inclusion with H. cequi-
costatus three types must be recognized. These types may be
called A, B and C to avoid confusion, but it must be understood
that they correspond with Miss Poole’s Types I, II & III.

Type A=Type I as defined by Gardiner. There is normally a
single small aperture on the basal surface in addition to the
opening of the Aspidosiphon chamber. In a few, this additional
aperture is absent, whilst in others, two or at most three such
apertures occur.

Type B is a much lower flattened form without a fossa; a
tendency for the coste to alternate; four septal cycles, the
exsertness of the septa of the fourth cycle on either side of the
primaries is very marked, and the joining over the tertiaries
and secondaries gives a characteristic star-like appearance ; the
columella is less well-developed, it is a compact trabecular mass,
and not distinctly papilliform as in Type A; the pores of the
endodermal canals are more numerous, and are not restricted to
the base, but are distributed irregularly round the lower part of
the corallum.

Type C is a taller form, the base tends to equal the disk in
diameter, and the walls are nearly perpendicular ; a fifth cycle of
the septa is present ; there is a distinct fossa, and the centrally
depressed columella is a trabeculate mass asin the previous type;
the pores of the endodermal canals are disposed in a ring a little
below the calicular margin.

In defining these three types, I have retained Gardiner’s Type 1
and: Miss Poole’s Type C in order to avoid confusion as far as
possible ; but it must be borne in mind that Type B as here
defined is very different from Gardiner’s Type IT.

Practically the whole of the South African collection belongs to
Type A, the Burmese collection belongs to Types B and C; the
specimens Miss Poole described as belonging to Type I should be
included in Type B. The Ceylon collection described by Professor
Bourne is somewhat intermediate between Types A and B. These
three types will embrace the species of Semper, Rehberg, and
Alcock*.

H. parasiticus Semper [38]. Intermediate between Types A

and B.
H. philippinensis Semper [38]. Two types. PI. xx. fig. 12 is
Type B. PI. xx. figs. 13 & 14, Type C.

* T have not considered H. sulcatus and H. lamellosus Verrill, and H. cochlea
Gmelin, as I have not been able to obtain first-hand reference to these species.

1026 MISS R. HARRISON AND PROF. S. J. HICKSON ON

H. pulchellus Rehberg [37]. Intermediate between Types B
and @. General facies, fifth septal cycle and depressed
columella, Type C. Alternating costze and distribution of
lateral pores, Type B.

H. oblongatus Rehberg [37]. Type C.

H. wood-masoni Alcock [1]. Type B.

The specimen from the Persian Gulf also belongs to Type B.

The other specimens of Heterocyathus in the Persian Gulf
collection constitute a new species. The tendency to remain
conical is very marked; the coste are distinct from those of other
species ; and the uniformity in size separate it sufficiently until a
good series of intermediates are found which will link it up
with /7. alternatus.

I therefore recognize in the genus the following species :-—

1. H. cequicostatus Milne-Edwards & Haime.
Stephanoseris rousseaui Milne-Kdwards & Haime.
H. parasiticus Semper.
H. philippinensis Semper.
H. oblongatus Rehberg.
H. pulchellus Rehberg.
H., wood-masoni Alcock.
2. H. alternatus Verrill.
H. cwequicostatus (Gardiner’s Type I).
3. H. heterocostatus, sp. 0.

Hrrerocyatuus aquicostatus Milne-Edwards & Haime [29].

A single specimen belonging to Type B. Coste, equal in
number to septa, extend to base, beset with irregular spines.
Base roughly granular. Slight calicular fossa. Septa in six
systems of four complete cycles, very spiny ; those of the fourth
eycle fuse with those of the third, and these in turn fuse with
those of the second. Inner margins of septa pass imperceptibly
into the trabecular columella.

Locality. Telegraph Cable, Persian Gulf. Depth 40 fathoms

HETEROCYATHUS HETEROCOSTATUS, sp. n. (Pl. LVII. fig. 6;
Pl. LVIII. fig. 14.)

Corallum simple, free, variable in shape from a low discoid
form to a conical cornuate form. Upper part of corallum
deeper in colour than base. Height varies between 3mm. and
7 mm., and calice between 3°53 mm. and 6 x 5mm. Coste of
two distinct types. Those corresponding to the septa of the first,
second and third cycles are visible from the base to the tip of the
calice, prominent sharp ridges with a single series of coarse
eranulations ; the alternate coste, corresponding to the septa of
the fourth cycle, extend only about half-way down the corallite
from the lip of the calice; they are less prominent, and beset
with numerous irregular granulations. Calice slightly elliptical,

CORALS FROM THE PERSIAN GULF. 1027

open, deep. Septa in four complete cycles. Primaries large, very
prominently exsert ; quaternaries join over the tertiaries, and again
deep down in the calice over the secondaries; quaternaries on either
side of the primaries much developed, and prominently exsert in
the manner typical of the genus Heterocyathus ; all septa beset with
numerous fine granulations arranged in radial ridges. Pali in the
form of small denticulations, which pass imperceptibly into ¢
parietal, fasciculate, centrally depressed columella. At the base, a
circular aperture leads to an Aspidosiphon chamber containing ¢
small Gastropod shell, in which a Sipunculid lives commensally
with the coral zoophyte.
Nine specimens.

Locality. Karachi. Depth 15-40 fms.; bottom, shell-sand.

2

=

Paracyatuus cavarus Alcock [1]. (Pl. LVII. fig. 5.)

Corallum simple, fixed by spreading base and expanding slightly
to lip of calice. Height varies between 15 mm. and 20 mm., but
the shortest has the largest calice. Calice variable in shape,
probably owing to local surroundings ; in some specimens calice is
circular, in others oval, while others have various indentations
and exerescences. Diameters of calices 15 x 15 mm., 16 x 12 mm.,
17x 10mm.,18x15mm., 20x13mm.,20x18mm. Coste visible
from base upwards, low broad ridges covered with minute
granulations, corresponding to each septum; towards the lip of
the calice these costal ridges become more marked, and somewhat
stouter and more exsert coste alternate with slightly smaller
coste. This alternation of larger and smaller costze is apparent
in some specimens at the extreme base of the corallum as well as
at the ealicular margin. Septa close, not markedly exsert ; the
size of the septa and the extent to which they are exsert
diminish in a descending order of magnitude according to the
eyele to which they belong: those of the first cycle are larger
than those of the second, the second than the third, and go on.
They are uniformly covered with minute granulations, which are
arranged in a series of longitudinal and radial rows. he calice
is widely open, and the inner margins of the septa are cut up
into large irregular nodules representing small pali, which pass
imperceptibly into a fasciculate parietal, centrally depressed
columella, ‘There appear to be typically five orders of septa, but
those of the lower orders are not always easy to interpret. ‘The
primaries are always large and extend to the columella; their
paliform nodules are slightly larger and more projecting than those
of the septa of lower orders; the septa of lower orders tend to fuse
together near the columella, and pass into it as an ill-defined mass
of minute pali. The larger size of the primary septa and the
grouping together of those of other orders give a hexagonal
starlike effect to the calice. In none of the seven specimens from
the Persian Gulf are the five cycles complete, although in one
there are 94 septa, only two septa of the fifth cycle being missing.

In the specimen figured (text-fig. 217), however, there are also

1028 MISS R. HARRISON AND PROF. 8S. J. HICKSON ON

94 septa, but here the arrangement is not so simple: in one lateral
chamber on each side between the primary and secondary septa
the fifth cycle is unrepresented, there being only three instead of
the normal seven septa. On the other hand, in both the apical

Text-fig. 217.

Diazram ui tic plan of the sah? of Paracyathus cavatus. The primary and
secondary septa are black, the tertiaries are cross-hatched, the quaternaries are
crossed with diagonal lines, the quinaries are left blank; i, ii, ii, iv, v, septa
of the first, second, third, fourth, and fifth cycles.

chambers at one end of the long axis there is an excess of septa.
In both chambers, between the primary and secondary septa on
the one side, and the primary and secondary septa on the other

CORALS FROM THE PERSIAN GULF. 1029

side, there are more than the normal number of septa. It
appears as if the tertiary septa of both these chambers had been
split by a wholly superfluous quaternary septum, while in one
case a quaternary septum has been further split by a super-
numerary quinary. The same phenomenon is apparent to a
greater or lesser degree in all the remaining five specimens. : In
one specimen there is a sudden outpushing of the wall of the
calice between a primary and a tertiary septum, in which there
are no fewer than nine instead of the normal three septa; in
every case there is a costa to correspond with each septum, and
the alternation of large and small septa and coste is maintained.

Seven specimens fixed on to a mass of mud, shell and serpulid
tubes.

Locality. From Cable 60 miles S.W. of Bushire, Persian Gulf,
Depth 30 fms.

Two specimens are infested by the Cirriped Pyrgoma stokesii,
which forms a bulbous gall within the wall of the coral (Pl.
LVII. fig. 5c). This distortion does not, however, interrupt the
regularity of the coste, which are distinctly visible on the outside
of the parasitic chamber; nor does the presence of this parasite
appear to affect the number of septa; there is no excess or
shortage of septa in the attacked specimens beyond that which is
normally observed in the unattacked individuals.

Genus Trematorrocuus T. Woods [39].

TREMATOTROCHUS ZELANDIZ. (Pl. LVII. fig. 4; Pl. LVIII.
figs. 15-17).

Conocyathus zelandie Duncan [14].

Corallum regularly conico-cylindrical, free, without trace of
attachment. No epitheca. Height of largest specimen 7 mm.,
‘diameter of calice 3°4 mm. Coste in four complete cycles,
‘prominent, smooth, and equal in the upper part of the corallum.
Only those of the first two cycles extend to the base; those of
the third cycle extend downwards for about three-quarters, and
those of the fourth cycle for a distance varying between a third
and three-fifths of the height of the whole corallum. The cost
between those of the second and third cycles are longer than the
costee between those of the first and third cycles; they do not
join those of the preceding cycle, but there is a thickening of the
costz of the first three cycles below the point at which that of
the succeeding cycle ends. Intercostal furrows penetrated by
minute, regularly disposed perforations (Pl. LVII. fig. 4). Calice
circular, no fossa, Septa in six systems of three complete cycles,
all exsert, those of the first cycle more prominently so than those
of the succeeding cycles; very thin and beset with small spinous
granulations. Septa of the third cycle join those of the second
a short distance below the lip of the calice ; they are incomplete
at their inner margins, large fenestrations occurring at the point
Proc. Zoot. Soc.—1911, No. LXX. 70

1030 MISS R. HARRISON AND PROF. S. J. HICKSON ON

where the septa of the third cycle join those of the second, and
those of the first and second join the columella. The lower
fifth of the corallum has been filled up internally by a secondary
deposit of calcareous matter, but the outline of the original struc-
ture is faintly discernible in photographs of sections at this level,
and reveals that the columella is parietal, formed by the union of
the inner ends of the septa.

Three specimens.

Locality. Persian Gulf. Depth not recorded.

The genus 7rematotrochus was proposed by Tenison Woods [39 |
for a fossil from the Miocene of Australia. The coral had all
the characters and appearance of a Turbinoliid, but with the
important difference that the wall was penetiated by large per-
forations between the coste, giving free communication between
the interseptal chambers and the exterior.

This remarkable coral did not receive the attention it deserved,
and subsequent authors received sceptically the statement that
these perforations were really present; for the point called in
question the validity of the division of the Madreporaria into
Perforata and Imperforata. Duncan [15] pointed out the diffi-
culty of placing this genus, but referred it to the Turbinoliide
in close relation with Turbinolia, Stylocyathus, Conocyathus, and
Bistylia*.

- In a series of papers published in the Transactions of the
Royal Society of South Australia, Dennant [7 & 8] has since
described six more fossil forms and one recent form (7. verconis)
from Australia and Australian Seas. In the first of these
papers he describes two species from Eocene and one species
from Miocene deposits. The recent species was found in St.
Vincent’s Gulf and Backstairs Passage at depths of from 15 to
22 fathoms. All have the characteristic perforations, which
fact caused him to remove the genus altogether from the Turbi-
noliide and place it among the Hupsammiide. Later he described
three different species from older Eocene beds than those in
which his previous species were found, and in two of these
the perforations do not pierce the wall, but are merely pore-
like cavities extending half-way through the thickness of
the wall like the intercostal dimples of some Turbinolide.
This made him put the genus back among the Turbinolide,
and he referred to Gregory’s [23] suggestion that the Perforate
type of coral has been derived from the Imperforate. He
further remarked on the wide distribution through time of the
original species 7’. fenestratus, and appended a drawing of a
portion of the wall cf one of them, placing the existence of the
pores beyond all suspicion of doubt. All the previously described
species come from Australia, and Dennant pointed out the

* Duncan uses the phrase “one of the species”’ as if more than one species were
known. At the time ofthe publication of his paper (1885) I am not aware that
any other species had been cescribed.

CORALS FROM THE PERSIAN GULF. 1031

occurrence of a recent form as evidence of the close relationship
between the living and fossil fauna ef that country. He made
no attempt to discuss the significance of these forms in con-
nection with the existing classification of the Madreporaria, but
dismissed the subject with the single statement—“ the broad
distinction usually made between perforate and non-perforate
corals breaks down.”

The occurrence of another recent species of this problem-
atical genus again forces the subject on the attention of the
systematist. That it isa T'rematotrochus is, | think, unquestion-
able. It agrees with Dennant’s latest definition in all points
but one :—He has described the septa as being solid; but in
T. zelandie the septa are incomplete, large fenestrations occurring
at the point at which the tertiaries join the secondaries, and the
primaries and secondaries join the false columella. Realizing
that the existence of these fenestrations and of the pores in the
outer wall might again be doubted (having already admitted
that these latter are minute and difficult to see), I have ground
sections of one specimen, thinking that actual photographs would
be more convincing than external observations. These sections
have been made at intervals of #, mm. and the entire coral has
been ground away; every section was photographed at a magni
fication of 20, and from these photographs a wax model has
been constructed. There is, therefore, a permanent record of the
structure of this most interesting coral, and it would be idle to
deny the existence of either thecal pores or septal fenestrations.
But in spite of this, I think that it is rightly placed among the
Imperforate corals.

In text-fig. 218, A and C are diagrammatic representations
of transverse sections of portions of the wall of a Heterocyathus
and a Heterepsammia (after Bourne [6'), for comparison with B,
a transverse section of 7. zelandiw. The Heterocyathus has a
solid theea embedded in stereoplasm. The Heteropsammia has
a porous spengy wall. A glance at the WVrematotrochus will
show that the extremely thin wall is incomplete, and that here
and there are small communications between the interseptal
chambers and the exterior; but it could not be regarded as a
“porous coenenchyme” such as is diagnostic of the division
Perforata. ‘The condition is much more that of a Fungiid which
has grown conically instead of spreading out with a widely open
oral surface, with that part of the wall between the perforations
representing synapticula. In some of the fossil forms the per-
forations are so large and the intervening solid structure little
more than a narrow bar, that the comparison with synapticula
is more striking than in the present instance. The analogy,
however, must not be pressed too far, but it widens the gulf
between 7rematotrochus and the Perforata, and tends to justify
its inclusion among the Imperforata.

_ Another point to be emphasized is the localities in which the
species are found. Hitherto the eight species of the genus were

CAO}

MISS R, HARRISON AND PROF. S. J. HICKSON ON

1032

ted area of Australia and the seas round

imi

all found in the |

the

in

but now we find it

7

that continent and New Zealand

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*(aumog «aqye) ru2payorm vrmunsdowazazy JO WN]]R109 ayy YSUOAYY UOTZOOS OLYBULLUBASLIP B JO WOO “{)

‘DIPUNIAZ SNYIOAZOINUALT, JO VANI[BAOD OY} YSNOAYy WOTZOIS O1yVUUUMIBISVIG “q

‘(ananog aeqye) sngnzsooinba snyjvhoowazezy JO WIN][R109 AY} YSNOAY} UOTZaS OTAVLUUTUBALSEIp B JO UOIWO “W

"BIG “8Y-489,

10

lentity of the last with Conocyathus zelandi

1€

The

Persian Gulf.

has already been pointed out in the introductory remarks of this

CORALS FROM THE PERSIAN GULF. 1033

paper, and the occurrence of the same species in localities so far
apart 1s remarkable.

I had hoped that the sections might throw some further
light on the question of septal sequence, but unfortunately the
lower fifth of the coral has been filled up solidly by a secondary
deposit of calcareous matter, and the interseptal chambers do
not begin to appear distinctly until a level is reached where
twelve septa and twenty-four coste are already present. If it

Text-fig. 219.

eae

A. Diagrammatic plan of the cost of Tremtotrochus zelandie, showing the
difference in length of the cost of the fourth cycle. i, ii, ili, iv, costae of the
first, second, third, and fourth cycles.

B. Diagrammatic plan of the septa of Ti-ematotrochus zelantie, representing the
exsert peripheral portion of the primary septa, and the exsert pali of the
secondary septa. i, li, iii, septa of the first, second, and third cycles; ea.s., exsert
portion of the septa of the first cycle; p., pall.

may be taken that the cost correspond to the septa and that the
base of a coral represents the earliest formation, it might be
objected that Pourtalés’ law for septal sequence has not been
followed in this case; for the cost which extend to the base,
twelve in number, are those corresponding to the first and
third (according to Pourtalés [35]) cycles of septa. That is to
say, if the secondary septa split peripherally in a Y-shape, and
a tertiary septum grew up between the arms of the Y, the
septa generally spoken of as secondary and tertiary ave named

1034 MISS R. HARRISON AND PROF. S. J. HICKSON ON

conversely to the order in which they appear in ontogeny. But
it must be remembered that the stem of the Y is also the
representative of the secondary septum according to either the
popular or Pourtalés’ nomenclature ; and these septa still exist
in their normal position midway between the primary septa,
and project upwards as pali, or paliform lobes: in the lower
portion of the colony, before the secondary septa have begun
to branch, the coste corresponding to them are also unbranched,
and extend to the base. A comparison of the diagrams A, B,
in text-fig. 219, illustrates this point.

Diagram A represents 4 plan of the cost and Ba plan of the
septa; in Aa line “aa” has been drawn across the lowest part
of the corallum, and in B a corresponding circle “aa” has been
drawn round the inner part of the septal plan; in both, the
costee and septa respectively of the first and second cycles are
present; presumably the coral must at one time have passed
through a stage when these cycles only were present. In
T. fenestratus this is the arrangement in the adult. Again,
another line “66” is drawn across A at a higher level and a
corresponding circle “bb” on B; in both, the cost and septa of
three cycles are present. When the secondary septa branched
peripherally, new costs arose in connexion with these branches,
and the original secondary eoste remained in connexion with
the inner unbranched part of the secondary septa which persist
as pali or paliform lobes. Bourne [6, text-fig. I. 2] has given
a coniprehensive diagram iHlustrating the relation of septa and
pali according to Pourtalés’ primciple: diagram B illustrates the
arrangement in 1. zelandie, which is entirely in agreement
with it.

The fact that the costz of the fourth cycle are of different lengths
(see text-fig. 219 A, iv) is in accordance with Duerden’s [11]
account of the septal sequence of Siderastrwa radians, in which
he states that new mesenterial pairs appear in some interseptal
chambers, before those in others of the same eycle.

AGELECYATHUS PERSICUS Duncan [14].

Corallites rising from an encrusting base, expanding slightly
from base to calice. Calice elliptical, depressed at one end of
the long axis. Height of corallum from 5-10 mm. at depressed
end of the long axis, and from 15-20 mm. at the opposite end.
Diameter of calice 9x75 mm. to 11x9 mm. The shape of the
corallites suggests that the colony was fixed on a vertical sur-
face, and the inequality of height in the two extremities of the
long axis is a response to an effort on the part. of the indi-
vidual corallites to assume a vertical position. Coste in four
complete cycles, distinct from base upwards, slightly exsert and
finely granular. Septa in six systems of four cycles of which
the last is incomplete, although all the coste of the fourth
cycle are present; septa of the first two cycles more exsert
than those of the third cycle; all three cycles reach the columella ;

CORALS FROM THE PERSIAN GULF. 1035

septa beset with minute granulations arranged in radial ridges.
An incomplete single crown of pali before the septa of the third
cycle. Columella fascicular.

Text-fig. 220.

Diagrammatic plan of the septaof Agelecyathus persicus. Septal orders represented
as in text-fic. 217. Six detached pali are represented. i. ii, iii, iv, septa of the
first, second, third and fourth cycles; p., pali.

A single specimen consisting of five individual corallites arising
from a common encrusting base.

Locality. From Cable 60 miles 8.W. of Bushire, Persian Gulf.
Depth 30 fms.

Before some of the septa of the third cycle there are distinct
upwardly projecting lobes corresponding to pali. Thus in the
calice figured (text-fig. 220) there are six such pali opposite six
septa of the third cycle, three at each end of the long axis;
there are two less conspicuous lobes before two septa of this
cycle, while the remaining four septa of the third cycle have
no trace of any such Jobe. Milne-Edwards and Haime [29]
divided the family Turbinoliide into two sub-families—the Tur-
binolinz without pali, and the Caryophylline with pali; and this
classification has found more favour with subsequent authors than
Dunecan’s [15] later classification, which divided the family into
Turbinolide simplices, Turbinolide gemmantes, and T'urbinolide
reptantes, according to their habits of growth and reproduction.
This coral affords evidence in favour of Dunean’s classification ;
indeed it would be difficult to know in which of Milne-Edwards’

1036 MISS R. HARRISON AND PROF. 8. J. HICKSON ON

and Haime’s groups to place it, for here in a single calice is a
combination of the diagnostic characters of the two sub-families :
pali are present before some and not before other septa of the
same cycle.

This specimen is considerably larger than Duncan's original
example from the same locality, and differs from his flew of
the type specimen in having the septa in the first two cycles
much exsert; but he calls attention to the variability of the
pali, the size and the number of the septa, as being features
of the individual corallites. The similarity in other details 3s
close.

Family FUNG611D &.

FuNG6IA PATELLA Ellis & Solander [16].

Three specimens from the Persian Gulf which are identical
with Doderlein’s figure [10, pl. i.] of the Cycloseris-form of
ff, patella, Wayland Vaughan’s figure [40, pls. xxv. & xxviii.] of
I’. patella, and Gardinexr’s figure [18, pl. 11x.] of Cycloseris hexa-
gonalis. They afford further evidence of the wide distribution
of the species, and for the justification of absorbing the genus
Cycloseris in the genus Fungia.

Locality. Twe specimens from Shaikh Shuaib Island, Persian
Gulf. Bottom, rock. Depth 10 fms. Exact locality of third
specimen not recorded.

Family HuPSAMMIID&.
HereroPsAMMIA APHRODES Alcock [1].

Eight specimens each with a single calice, The inflated spongy
exsert edges of the septa of the first two cycles are well developed.
Locality. Telegraph Cable, Persian Gult. Depth 49 fms.

DENDROPHYLLIA sp. ? de Blainville [5]. (Pl. LVIL. fig. 7.)

Colony bushy, springing from an encrusting base on which are
numerous small individuals. Budding lateral or basal. Polyps
occasionally joined by horizontal bars, but in every case such bars
are inhabited by a small commensal worm, and have probably
been formed in connexion with this parasite. Polyps tend to
assume a vertical position, but one polyp which appears to
arise from a horizontal bar is directed downwards. Size of the
polyps very variable; some searcely rising from the encrusting
base; some attaining to a height of 25 mm. Calice varies
between 2x2 mm. in the youngest to 7x6 mm. in the oldest
individuals. Coste distinct from base upwards, equal, granular,
not exsert, correspond in number to the septa. Septa in
six systems of four cycles with a few representatives of the
fifth cycle, all irregular. Some septa of the lower cycles are
grouped together round that of the preceding cycle in the typical
dendrophylliid manner, while others extend to the columella, and
are equal in size to those of preceding cycles. All septa are

CORALS FROM THE PERSIAN GULF. 1037

finely granular, the lower orders are perforate. Calicular fossa
very deep. Columella fascicular, parietal; well developed in older
individuals.

Locality. From Cable 60 miles 8.W. of Bushire, Persian Gulf.
Depth 30 fms.

With only a single specimen of a coral such as this, which
exhibits such a wide range of variability, the creation of
a new species has not seemed to me justifiable. I have, therefore,
merely noted its characters and appended a photograph of the
specimen, until such time as it may be found in greater numbers.

Note on Miss Harrison’s memoir on some Madreporaria from
the Persian Gulf, and some further notes on Pyrophylli«
inflata. By Sypney J. Hickson, M.A., F.R.S., F.Z.8.

The manuscript of Miss Harrison’s paper was sent to me shortly
before she left this country for India, with a request that I would
read it and revise it for publication. The number of species in
the collection is small and there is only one that is new to science,
but there are several points in the paper which it seemed to me
required rather fuller consideration than she has given to them,
and I have ventured therefore to write an addendum, leaving her
original memoir intact. Had she remained in England I would
have suggested the inclusion of these remarks in her paper, but
under the circumstances, I think it is better to publish them
under a separate title and thereby take the whole responsibility
for them. I have rearranged the order of the species, furnished
the list on p. 1019, and added the family names, but in other
respects the paper is as it left her hands.

At the present time our knowledge of the fauna of the Persian
Gulf is very limited. None of the great deep-sea exploring
expeditions have visited it, and independent investigators with
sutticient knowledge and energy in this region have been few and
far between.

Mr. F. W. Townsend, of the telegraph staff of the Indian
Government, bas made a large and valuable collection of shells,
and these have been described in a series of papers by Cosmo
Melvill and Standen,* but so far as I can discover, very few
genera and species belonging to other groups of marine animals
have been recorded. The richness and interest of the Molluscan
fauna suggests that many new forms have still to be discovered
in the Gulf, but the subject of special importance that the study of
this fauna would shed light upon, is the relation of the fauna
of the Persian Gulf to that of the Mediterranean Sea. It has
frequently been suggested that in the past there was a connexion
between the Indian Ocean and Atlantic Ocean by way of the
Mediterranean Sea ; and, judging by the present day geographical
features, it is probable that the last connexion between them

* Wor list of papers see Proc. Zool. Soc. 1906, p. 783.

1038 MISS R. HARRISON AND PROF. S. J. HICKSON ON

was broken by the formation of the Isthmus of Suez. But the
last connexion but one was that by way of a strait, of which
the remaining part is now the Persian Gulf; and this gulf with
its narrow outlet into the Indian Ocean, high temperature and
great rivers, might be expected to retain some of the fauna which
had been subject to very similar conditions in the South-east
corner of the Mediterranean Sea. Melvill and Standen call
attention to the considerable generic analogy between the Mollusca
of the two regions, and point out that the species of the
Persian Gulf show close affinities with South-Huropean forms.

It would be quite premature to draw any far-reaching
generalisations as to the distribution of Madreporaria from the
few specimens that are here recorded, but attention may be
called to one or two points of general interest.

The occurrence of Pyrophyllia in deep water in the Persian
Gulf is interesting from the point of view of geographical
distribution. Alcock [3] in his comments on the deep-sea Madre-
poraria of the Indian Ocean, calls attention to the ‘“ many
intimate aftinities of the fauna of moderate depths of the Indian
seas with the North Atlantic fauna,” and considers them ‘to be
sufficient to suggest a direct sea-connexion, in the past, between
the Atlantic and Indian Oceans, and the case of Caryophyllia
communis and Flabellum laciniatum would indicate that the
connexion was by way of the Mediterranean.”

The case of Pyrophyllia and Guynia appears to me to give
even stronger evidence of the truth of this hypothesis, than that of
the two species quoted. The genera Caryophyllia and Plabellum
are both very widely distributed recent corals, and it is possible
that in comparatively recent times these two species may have
had an almost cosmopolitan distribution. Guynia and Pyrophyllha
are, so far as is known at present, very restricted in their distri-
bution and are totally unlike any other recent coral—with the
possible exception of the West Indian genus Haplophyllia.

Although so much alike in important characters, they are
sufficiently distinct for us to believe that they were separated
from one another at a very remote period. It is rather more
difficult to believe that the Indian Ocean and Mediterranean
specimens of Caryophyllia communis and Flabellwm laciniatwm
can have undergone no differential change since the time when
the Mediterranean Sea and the Indian Ocean were in commu-
nication. A second point of interest is, that the only other coral
with which Pyrophyllia shows aflinities, namely Conosmila,
should be found in the Tertiary deposits of Australia. Standing
by itself, this is only one of those facts of geographical distribution
which it is important to note but impossible to explain in a
satisfactory way. But its importance as a fact is emphasized
when it is placed side by side with the facts of the distribution of
Trematotrochus.

As Miss Harrison points out, the specimens of Z’rematotrechus

CORALS FROM THE PERSIAN GULF. 1039

found in the Persian Gulf are closely related to species of corals
from the Tertiary deposits of Australia, and to a recent coral
found at depths of from 15 to 22 fathoms in St. Vineent Gulf,* and
ilentical with a recent coral from “no very great depth” im
Cook’s Strait, New Zealand.

Of the geograpical distribution of the other species very little need
be said. Heterocyathus heterocostatus is new, but a closely related
species H. equicostatus, of which one specimen was found, appears
to be widely distributed in the Indian Ocean. Pwracyathus cavatus
is found in the Indian Ocean and is said by Alcock to he closely
related to the Hocene fossil P. crassus from the London Clay.
Agelecyathus persicus was previously recorded by Duncan from
the Persian Gulf, but was said to oecur also off St. Helena.
Fungia patella occurs in the Indian Ocean. ‘The type specimen
of Flabellum magnificwm was found off Sumatra at a depth of
470 metres. The genera Heteropsamnia and Dendrophylliaappear
to be widely distributed in the Indian Ocean. So far then as
this smail collection of corals is concerned, Pyrophyllia inflata is
the only species that even suggests a former connexion of the
Gulf with the Mediterranean Sea.

The genus Pyrophyllia was briefly described in the ‘‘ Manchester
Memoirs,” 1910; but it may be convenient to take this
opportunity of adding a few general remarks on the genus and
of publishing some further illustrations (P]. LVII. figs. 8-11;
Pl. LVITTI. figs. 18, 19).

Pyrophyllia inflata is a small unattached solitary coral, of
about 4 to 5mm. in length, and 1 mm. in diameter at the margin
of the calyx.

The two most important characters are :—(1) Its very
pronounced and-invariable octoradiate symmetry, and (2) the
presence on the external surface of well-marked accretion ridges
with short but definite costal spines (text-fig. 221).

Pyrophyllia is related to the recent genus Guynia, and
to the extinct Tertiary genus Conosmilia. The number of
septa in Pyrophyllia and in Guynia is sixteen, and of these eight
are larger and may be called the*primary septa, and the remaining
eight are smaller and may be called the secondary septa.

In Guyma one of the eight primary septa is larger than
the others, this large septum being according to Duncan a
““very marked rugose peculiarity.” Moreover, in Guynia four of
the primary septa are sometimes larger than the other four, so
that according to Duncan the system of septa is four primary
septa, four secondary septa, and eight tertiary septa.

In these respects and in others of less importance, Guynia
seems to be distinct from Pyrophyllia. In the genus Conosmilia
there is a more variable arrangement of the septa. In
C. elegans, C. litwolus, and C. anomala there are eight
primary septa, eight secondary septa, and thirty-two tertiary

* The specimens of this coral Z. verconis were much worn and were no doubt
dead corals when collected.

1040 MISS R. HARRISON AND PROF. S. J. HICKSON ON

septa; but in C. striata there are only six primary septa, six
secondary septa, and twelve tertiary septa. The more recently
described species of this genus, C. granulata and C. stylifera
(Dennant 7), agree with C. striata in the hexaradial arrangement
of the septa. In Conosmilia, moreover, the accretion ridges do
not appear to exist unless they are represented by the “ beautiful
herring-bone ornamentation of the surface.”

Diagrammatic sketches of the septal arrangement of Pyrophyllia inflata.
g 12) g Yrop

c., columella; m.s., secondary septa; p.s., primary septa.

A. Arrangement of the septa just below the margin of the calyx. B. At the base
of the calyx. C. In the lower parts of the coral. (From Mem. Manch. Lit.
Phil. Soc. 1910.)

The columella of Conosmilia resembles Pyrophyllia im being
laminate and in this respect differs from Guynia, in which the
columella is cylindrical.

There has been a great deal of hesitation in giving the two genera
Guynia and Conosmilia a definite resting-place in the system of
corals,

Duncan * at first placed the genus Guynia in the Order
Rugosa and in the family Cyathaxoniide, but subsequently
removed it [15] to the family Turbinoliide. Miss Ogilvie placed
it in her new family Amphiastreeidze +.

Dunean $ at first placed the genus Conosmilia in the Order
Rugosa, family Stauride, but subsequently removed it to the family
Astreidee Simplices and placed it close to the genus Zrochosmilia.
Miss Ogilvie placed this genus in the family Turbinoliide.

In my preliminary account of the genus, I remarked that ‘it
cannot be denied that Pyrophyllia has some characters reminiscent
of the extinct Rugosa,” but on reconsideration I do not feel

* Phil. Trans. R. §. 1872. y Phil. Trans. 1896. £ Phil. Trans. 1872.

CORALS FROM THE PERSIAN GULF. -1041

satisfied that these characters—the octoradiate symmetry, the
pronounced accretion ridges and the septal fusions in the lower
part of the coral—really indicate any true affinities with Paleeozoic
corals. Duncan found one specimen of Guynia in which, owing
to the abortion (?) of two septa, the upper part of the corallum
showed a hexaradiate symmetry. This specimen was regarded
by him as ‘‘very suggestive in the matter of the evolution of
the hexameral from the octomeral types, or rather from the
tetrameral.” But the view that was in his mind, that the
modern hexaradiate corals are descended from a previous tetra-
radiate type, is not one which commends itself to more recent
investigators of coral structure. The researches of Duerden*
suggest that the tetraradiate symmetry of Rugose corals is derived
from a more primitive hexaradiate symmetry, and there is no
evidence in the development of modern corals, or satisfactory
evidence in paleontology, that the modern hexaradiate corals are
derived from a previous tetraradiate or octoradiate ancestry.

It is true that many Paleozoic corals exhibit four dominant
primary septa, but there are none with eight dominant septa.
The true octoradiate condition is known only in certain species of
the Tertiary genus Conosmilia and in the recent genera Guynia
and Pyrophyllia. ‘The recent genus Haplophillia has also eight
primary septa and eight secondary septa, but it is suggested by
Gardiner that this genus is only a growth stage of “Duncania.
The close relationship of Conosmilia to the simple modern
hexaradiate corals suggested by the classification of Duncan and
of Ogilvie, seems to point to the conclusion that the Guyniide
represent an early offshoot of the modern hexaradiate line of
descent, and that their true affinities lie rather with the modern
Turbinoliide than with any known Paleozoic coral.

I may refer in conclusion to some remarks made in my
preliminary paper on the presence of endotheca in Pyrophyllia.
Although Duncan’s definition of endotheca is not very explicit,
I have come to the conclusion that there is no structure in
Pyrophyllia that really corresponds with what is usually regarded
as endotheca by the paleontologists. A longitudinal section of a
Pyrophylia (Pl. LVITI. fig. 19) shows three regions: an upper
calicular region (a) in which the septa are free from one another
and from the columella, a middle region (y) in which the septa
are more or less irregularly fused with one another and with the
columella, and a lower region (z) in which the septa are again free
and the thecal wall inflated.

It seems probable from the texture of the well-preserved
specimens that the living tissues were confined to the upper
calicular region. ‘The growth in thickness and the fusion of septa
and columella in the middle region (see text-fig. 221) perform the
same function as the endotheca of Paleozoic corals, as they ‘“ unite
septa, close the loculi, and enable the coral to grow in height
and strength, and limit the growth downwards of mesenteries

* Ann. N. H. xviii. 1906.

1042 MISS R. HARRISON AND PROF. S. J. HICKSON ON

and soft parts,” but there are no thin plate-like structures
corresponding with those that constitute the greater part of the
endotheca of Paleozoic corals.

13.

14.

15.
16.
sf
18.

. Atcock, A.

. DENNANT, J.

BIBLIOGRAPHY.

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Indian Seas.” Journ. Asiatic Soc. Bengal, Ixil. p. 138.
Natural History Notes from H.M. Indian

7” Journ. Asiatic

Marine Survey Steamer ‘ Investigator.
Soc. Bengal, Ixii. p. 169.

. Aucock, A.—An Account of the Madreporaria collected by the

Royal Indian Survey Ship ‘ Investigator.’ Calcutta, 1898,
p. 29.

. Atcock, A.—‘“‘ Report on the Madreporaria of the Siboga

Expedition.” Siboga-Expeditie, Monogr. xvi. a, 1902, p. 52.

. DE Brarnvitiz, H. M. D.— Dict. des Sci. Nat. lx. 1830, p. 319.
. Bourne, G. C.—* Solitary Corals collected by Professor

Herdman in Ceylon, 1902.” Report of the Pearl Oyster
Fisheries in the Gulf of Manaar, iv. 1905, p. 187.

Dennant, J.—‘‘ Descriptions of New Species of Corals from
the Australian Tertiaries.” Trans. Roy. Soc. South
Australia, xxiii. pp. 112 & 228; xxv. 1901, p. 48.

“Recent Corals from the South Australian
and Victorian Coasts” Trans. R. 8. South Australia, xxvii.
1904, p. 1.

. Doperitetn, L.— ‘Die Korallen-Gattung Fungia.” Zool.
b) to}

Anz. xxiv. 1901, p. 353.

. Doperte, L.—‘‘ Die Korallen-Gattung Pungia.” Abh. v.

d. Senckenbergischen naturforschenden Gesellschaft, xxvil.

1905, p. l.

. Durrpen, J. E.—‘‘ The Morphology of the Madreporaria. V.

Septal Sequence.” Biol. Bull. vil. 1904, p. 79.

. Duncan, P. M.—‘‘On the Madreporaria of the Expedition of

H.M.S. ‘ Porcupine.” Proc. Roy. Soc. London, xviii. 1870,
e280:

Dean, P. M.—‘“ A Description of the Madreporaria dredged

up during the Expedition of H.M.S. ‘Porcupine.’” Trans.

Zool. Soc. London, viii. 1874, p. 303; x. 1879, p. 235.

Duncan, P. M.—“ Notices of some Deep-Sea and Littoral

Corals from the Atlantic Ocean, Caribbean, Indian, New-

Zealand, Persian Gulf and Japanese &c. Seas.” P. Z.S. 1876,
. 428.

inca P. M.—‘“ A Revision of the Madreporaria.” Journ.

Linn. Soe., Zoology, xviii. 1885, p. 1.

Exuis, J., & SOLANDER, D.—‘‘ Natural History of Zoophytes.”

1786.

GARDINER, J. 8.—‘‘On the Solitary Corals collected by the

author in the South Pacific.” P. Z.S. 1898, p. 525.

GARDINER, J. S.—“ On the Solitary Corals collected by Dr. A.

Willey.” Willey’s Zool. Results, 1899, p. 161.°

Ye),

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2l.

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24.

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35.
36.
37.
38.

CORALS FROM THE PERSIAN GULF. 1043

Garpiner, J. S.—“South African Corals, of the Genus
Flabellum.” Marine Investigations in 8. Africa, II., Cape

> Town, 1904, p. 117.

Garpiner, J. S.—“The Turbinolid Corals of 8. Africa.”

Marine Investigations in 8. Africa, III., Cape Town, 1904,
2 45,

Gasset J. S—“The Fauna and Geography of the

Maldive and lLaccadive Archipelagoes.” IT. Suppl. i.

Madreporaria, Pt. 111., Fungiide, Pt.iv., Turbinolide, p. 933.

GARDINER, J. S.—‘* The Percy Sladen Trust Expedition to

the Indian Ocean in 1905. The Madreporarian Corals.—I.

The Family Fungiide. Trans. Linn. Soc., Zoology, xii. 1909,
B25c

Lee J. W.— “Jurassic Fauna of Cutch. Corals.”

Paleonto]. Indica, 9th Ser. vol. 11., 1900.

Harrison, R. M., & Pootr, M.—‘‘ Marine Fauna from

Mergui Archipelago, Lower Burmah, collected by Jas. J.

Simpson and R. N. Rudmose-Brown :—Madreporaria.”

IP aise IOS), yon ele

Hickson, 8. J.—‘‘On a new Octoradiate Coral, Pyrophyllia

inflata.” Mem. & Proc. of the Manchester Lit. & Phil. Soc.

liv: Pt. ms, No, 12; 1910.

Kent, W. Savitte.—‘*On some new and little-known

Species of Madrepores, or Stony Corals, in the British

Museum Collection.” Proc. Zool. Soc. 1871, p. 285.

Lacaze-Dutuiers, H. pe.—‘‘ Faune du Golfe du Lion.’

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Ann. des Sci. Nat., 3° sér. ix. 1848, p. 211.

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Ann. des Sci. Nat., 3° sér. x. 1848, p. 65.

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Coralliaires.” 3 vols., Paris 1857.

Mosstey, H. N.—“ Deep-Sea Corals.” ‘Challenger’ Reports,

Zoology, 11. 1881, p. 127.

Ortmann, A. E.—“ Beobachtungen an Steinkorallen von der

Sudkiiste Ceylons.” Zool. Jahrb., Syst. iv. 1889 p. 493.

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Zoology, xvi. 1886, p. 203.

ReHBERG, H.—“* Neue und wenig bekannte Korallen.” Abh.

Ver. Hamburg, xi.

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f. wiss. Zool. xxii. 1872, p. 235.

1044 ON CORALS FROM THE PERSIAN GULF.

39.
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Trnison- Woops, J. E.—-‘On some Tertiary Fossil Corals and
Polyzoa.” Trans. Roy. Soc. New South Wales, xii. p. 57.
Vaucuan, T. W.—‘ A Critical Review of the Literature on
the Simple Genera of the Madreporaria Fungida, with a
tentative Classification.” Proc. U.S. Nat. Mus, xxviii.
1905, p. 371.

Vaueuan, T. W.—‘‘ Recent Madreporaria of the Hawaiian
Islands and Laysan.”” U.S. Nat. Mus. Bull. lix., 1907.
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Amer. Journ. Sci. & Arts, 2nd ser. xlix., 1870.

EXPLANATION OF THE PLATES.
Prate LVII.

1, Flabellam magnificum von Marenzeller. Viewed from above. x 3 diam.

2. Flabellum magnificum. Viewed from below to show the tubular rootlets of
attachment.

3. Flabellum magnificum. Side view.

4. Trematotrochus zelandie Duncan. Transverse section of the coral.

x ca. 12 diam. Compare text-fig. 218 3, p. 1032.

5. Paracyathus cavatus Alcock. A. View of avertical fracture. B. View from
above. C. Side view showing on both sides above, galls formed by Pyrgoma.
Nat. size.

6. Hetcrocyathus heterocostatus, sp.n. Nine specimens showing the variations
in height. x 2 diam.

7. Dendrophyllia sp.? x % diam.

8. Pyrophyllia inflata. Side view. x 5 diam.

9. Pyrophyllia inflata. View of the mouth of the calyx showing the columella

and primary septa.

10. Pyrophyllia inflata. View of a specimen that has been broken in half
longitudinally showing the irregular arrangement of the fused septa in
the lower regions of the ecral.

11. Pyrophyllia inflata. View of a specimen showing the fusion of the eight
primary septa and the secondary septa. Compare text-fig. 221 B, p. 1040

Prare LVIII.

ig. 12. Heterocyathus equicostatus M.-E. & H. Two adjacent costee which are

low broad ridges covered all over with uniform fine granules, the
intercostal furrows being small and shallow. ‘his speéimen belcngs to
Gardiner’s Type I.

13. Heterocyathus alternatus Verrill. Two adjacent cost. The costs on
the right (A) is the type of costa corresponding with the septal cycles
i, ii, andiii. The costa on the left corresponds to the septal cycle iv.
The intercostal furrows are deep. This specimen belongs to Gardiner’s
H. e@quicostatus Type 1.

14. Heterocyathus heterocostatus, sp.n. Side view. x ca. 14 diam.

15. Trematotrochus zelandie Duncan. Side view. x ca. 12 diam. Compare
text-fig. 219 A, p. 1033.

16. Trematotrochus zelandie. View of the base.

™ Trematotrochus zelandie. View of the calyx from above. Compare text-

fig. 219 B.

18. Pyrophyllia infiata Hickson. View of the calyx showing the columella,
the eight primary septa, four of the eight secondary septa, and the limes of
accretion.

19. Pyrophyllia inflata. Wongitudinal vertical section showing the fusion of
the primary septa in the lower parts of the coral. «, region of the calyx ;
y, middle region; z, inflated base.

‘P.

pith etiam

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SMC E 12

A

ON VARIATION IN A JELLYFISH. 1045

47. On Variation in the Medusa of Merisia lyonsi. By
Cartes L. Boutuncer, M.A., F.Z.S., Lecturer on
Zoology in the University of Birmingham.

[Received May 23, 1911: Read June 27, 1911.]
(Plate LIX.* & Text-figures 222-228.)

In 1908 I published an account [1]+ of a new lacustrine
Hydromedusan, Marisia lyousi, obtained by Dr. Cunnington and
myself from the brackish waters of Lake Qurun in the Fayim
Province of Egypt.

In my paper, whilst describing the anatomy of this interesting
form, | called attention to the fact that the number of tentacles
and radial canals in the medusa stage was subject to much
variation, and mentioned that in a series of 400 individuals
which I examined, 55, or nearly 14 per cent., differed from the
normal.

My description of the abnormal specimens was very short,
and it has been suggested to me that it would be of interest to
furnish a more detailed account of the variation of this medusa,
as well as to figure some of the more peculiar abnormalities.
This I was all the more prepared to do, as further study of the
collection had revealed additional points of interest in connection
with this phenomenon.

Variation is known to occur frequently in jelly-fishes ¢, and in
some species it has been very carefully studied ; although in many
cases the series dealt with were numerically far greater than the
one at my disposal, I know of no form in which such a variety of
abnormalities occur as in Merisia. The interest of the series I
am about to describe is not diminished by the fact that all its
members were collected in one locality and belong to the same
sex.

As the greater part of my material had already been distributed
when I decided to take up this subject again, it was necessary to
re-examine the specimens in the Natural History Museum,
London, and in the Cambridge Museum of Zoology; for permis-
sion to do this, I have te thank Mr. R. Kirkpatrick and Mr. L.
Doncaster, under whose charge the specimens are preserved in
these institutions.

In my description of Marisia lyonsi, I drew up the following
table to show the number and arrangement of the radial canals

* For explanation of the Plate see p. 1056.
+ The figures in brackets refer to the List of References on p. 1055.
t Cf. List of References on p. 1055.

Proc. Zoou. Soc.—1911, No. LX XI, 71

1046 MR. C. L. BOULENGER ON
and tentacles in 400 meduse taken at random from the material
at my disposal :—

Table showing the Number and Arrangement of the Radial Canals
and Tentacles in 400 individuals.

Number of Tentacles.
Sumter |Syeet of} ——— :

Individuals. Canals. | Perradial. ‘Interradial.|Adradial.! Subradial. | Total.

1 3 3 = = — 3

345 4. 4 — — — 4,

1 4 4 1 — _ 5

1 4 4 1 2 = 7

i 4 4, = 1 — 5

2 4 4 2 4 — 10

10 4 4 4 = = 8

28 4 4 4 8 =m | 16

1 4, 4 4 8 6 | 22

5 5 — — — | 6

1 6 6 — — = 6

Total 400

It will be noticed that the 55 abnormal individuals fall naturally
into two well-marked groups :—(@) which includes those medusze
which deviate from the normal tetramerous symmetry ; and (8),
which includes medusz with the normal number of radial canals
and primary, perradial tentacles, but possessing, in addition,
secondary tentacles situated between the four primary ones and
not connected with the stomach by means of radial canals.

It is my intention to discuss these two groups separately, as
it is obvious that in them we are dealing with two completely
distinct phenomena.

(a) The normal medusa of Merisca lyonsi is provided with
four radial canals at the distal extremities of which the four
perradial tentacles are given off. The gonad-bearing region of
the stomach is produced into four perradially situated pouches
which in the adults extend as finger-shaped diverticula for a
considerable distance along the radial canals.

Eleven specimens out of the 400 medusze examined (7. e. 2°75
percent.) were found to deviate from this general tetramerous
symmetry ; one medusa possessed only three radial canals, three
tentacles and three gonadial diverticula ; nine meduse had five, and
one had six of these structures. This type of variation is known
to occur in many species of meduse which are normally tetra-
merous in symmetry ; among craspedote forms it has been studied
best in Obelia (Hucope) [2], Clytia [3], Sarsia [4, 5], Rathkea
(Lizzia) [7], Podocoryne [8], Gonionemus [8], and Limnocnida [9].

VARIATION IN A JELLYFISH. 1047

Limnocnida being another African lacustrine form is of
particular interest. Giinther found that out of 70 individuals
collected by Dr. Cunnington in Lake Tanganyika, 54 meduse
showed the typical number (four) of radial canals, whilst 16, or
24 per cent., had five or more ; as in the case of Marisia, specimens
with five radial canals were commoner than those with a larger
number. Ginther, however, found no meduse with only three
canals, but such trimerous forms have been described in other

genera, e.g. Podocoryne [8| and Rathkea |7].

Text-fig. 222.

Merisia lyonsi.

Diagrams to show the arrangement of the tentacles im four medusze.

B represeuts the normal tetramerous form.

Similar variations occur in several members of the Scypho-
medusz: perhaps the most complete account of the phenomenon is
that given by KE. T. Browne for Aurelia aurita [11, 12]. This
author examined the variation in the number of tentaculocysts

in the ephyra larve and adults of the jelly-fish, and found that
713

1048 MR. C. L. BOULENGER ON

20°9°/, and 22°/,, respectively, were abnormal, the number of
these sense-organs ranging between 6 and 15, the normal number
being, of course, 8. ‘These figures, however, include a number of
specimens which are undoubtedly teratological monstrosities.

Browne pointed out that variation in the number of tentaculo-
cysts does not necessarily interfere with the other organs of the
body, but that there is a correlated variation between the number of
genital pouches and buccal arms as shewn by eight specimens :—
4 individuals had 3 genital pouches and 3 buccal arms, 1 had 5
pouches and 5 arms, and 3 had 6 of these organs.

In Merisia there must necessarily be a correlation between
the number of primary tentacles and radial canals, since recent
researches on the development of the gonophores of Hydromedusze
[13, 14] have shown that these structures arise together from the
endodermal pouches of the young medusa-bud,

There is also a correlated variation between the number of
radial canals and gonadial pouches, and this is only what we
should expect as the latter structures spread outwards from the
stomach on to the radial canals. ‘The single 6-rayed specimen,
however, had one gonadial pouch very poorly developed although
the other five were quite normal.

It is interesting to note that variation in the number of
primary radii in the medusa does not necessarily affect the
symmetry of the manubrium; this point is well shown in the
sections figured on PI. LIX.

A normal medusa with four tentacles, radial canals and
gonadial diverticula has the distal part of the manubrium
provided with a similar number of conspicuous endodermal
ridges or teniole, which can be seen in the section figured
Pl. LIX. fig. 1); a similar section through a pentamerous
individual (Pl. LIX. fig. 2) demonstrates the fact that, although
five canals and gonads are well developed and the manubrium
appears almost pentagonal, the number of endodermal ridges is
not affected, although their symmetrical arrangement is disturbed.

The above is in accordance with the observations of other
investigators :—Agassiz and Woodworth [2] examined 4000
specimens of Obelia (Hucope), but noted no variations in the
shape of the digestive cavity or in the number of the actinal lobes
of the manubrium, even in specimens with five or six radial
canals in place of the normal number (four) the actinal lobes
being always found to be four in number.

A. G. Mayer [3] has also made some interesting observations
which bear on this point. This author has made a careful study
of the medusa Pseudoclytia pentata, the only Leptomedusan
which is normally pentamerous, and which he considers to be
derived from some species of Clytia (Hpenthis), e.g. C. folleata.
In Pseudoelytia pentata there is much variation both in the number
of radial canals and in the number of oral lips; but whilst the
former incline towards the production of more than five canals,
the oral lips show a decided tendency to revert to the ancestral
condition of four.

VARIATION IN A JELLYFISH, 1049

As mentioned before, variation in the nwouber of radii is
of common occurrence in meduse, and in the above-described
varieties of Merisia lyonsi we are undoubtedly dealing with
a quite ordinary case of meristic variation. As Bateson remarks
in his well-known book on the Study of Variation [15] :—
“In radial series phenomena analogous to those of the variation
in linear series are seen in their simplest form. Just as in linear
series the number of members may be changed by a reconstitution
of the whole series so that it is impossible to point to any one
member as the one lost or added, so may it be in the meristic
variation of radial series: and again as in linear series, single
members of the series may divide. Between these there is no
clear line of distinction.”

In the abnormal specimens of Mwrisia, I do not think that we
are dealing with cases of division or suppression of one or more
radi ; from what we now know of the development of medusa-buds
[13, 14], it seems that the variation is in the number of radial
pouches formed in the early bud.

Since each pouch develops the radial canal and perradial tentacle
of its own section of the medusa-bell, such variation explains the
correlation of the numbers of these two sets of organs. The fact
that the manubrium is independent of the organs situated in the
umbrella is a point decidedly in favour of this view. I have,
unfortunately, not been able to study the development of an
abnormal specimen, all the medusa-buds I have sectioned proved
to be of normal tetramerous symmetry.

(>) The second group of abnormal individuals includes a
number of tetramerous medusze which bear secondary tentacles
between the four primary perradial ones. These secondary
tentacles may be interradial, adradial or subradial in position,
and differ from the perradial ones in not communicating with the
gastric cavity by means of radial canals; they are developed from
the ectodermal and endodermal cells of the umbrella-margin, their
cavities being in communication with the circular canal.

Altogether 44 of the 400 tabulated individuals exhibited this
kind of “abnormality, which is of a very peculiar type ; I have not
been able to find any record of a similar case in a medusa which
normally bears primary tentacles only. There are, of course,
numerous tetramerous meduse which normally posssess. such
secondary tentacles (e. g., Podocoryne, Turritopsis, Oceania,
Limnocnida), and they are known to start life with the fours
primary ones only, the secondary tentacles being developed as
the animals grow, usually in some definite sequence. Thus, in
the majority of cases, the second set of tentacles to appear is the
interradial one, four of these structures growing simultaneously
from the umbrella-margin between the four perradial tentacles.
Hight adradial tentacles are formed next, two in each quadrant
occupying the interspaces between the interradial and perradial
ones, and their formation may be followed by the appearance of a
varying number of subvadial tentacles.

1050 MR. C. L. BOULENGER ON

In such medusz it is obvious that the number of secondary
tentacles is correlated with the size of the individuals, the older
and larger meduse possessing a greater number of these structures
than the smaller and younger ones. In Merisia we are certainly

Text-fig. 223.

Medusa bearing four interradial tentacles im addition te the fenr perradial ones at
the extremities of the radial canals. In this and in the three following text-
figures A represents the medusa as seen in a side view, B its oral aspect.

Text-fig. 224,

Medusa with four interradial and eight adradial tentacles in addition
to the four primary ones.

not dealing with a phenomenon of this kind, for the larger and
more mature meduse are, with rare exceptions, unprovided with
secondary tentacles. In order to show this point more clearly,
I divided the 400 individuals which I examined into two

VARIATION IN A JELLYFISH, 1051

groups :—(i.) including meduse with an umbrella-diameter
measuring between 3 and 2 mm., and (ii.) with a diameter
measuring from 24 mm upwards,

Group (i.) contained 278 individuals of which 39, or about
14 °/,, possessed supernumerary tentacles between the four
primary ones, whereas of the 122 larger medusze belonging to
group (ii.) only 5 were provided with such structures.

Sexual dimorphism is known to occur in a few species of
medusze,* but I have assured myself that this is not the case
with the specimens I am describing; microscopic examination,
by means of sections and whole mounts, showed that all the
meduse’ of Merisia in my collection belonged to the male sex,
whether bearing secondary tentacles or not.

It must be obvious from the above account that the kind of
aviation I have just described is of a nature totally different from
that treated of in the first section of this note. We are certainly
not dealing with a case of ordinary meristic variation, and the
only conclusion I can arrive at is that these multitentacular
meduse form a distinct variety which has arisen as a mutation
from the normal form : it will be interesting to discover whether
this variety will maintain itself in Lake Qurun.

Altogether, forty-six f multitentacular specimens have been
examined by me; it was found that the number of secondary
tentacles varied considerably. 10 individuals possessed four
interradial tentacles as well as the four primary perradial ones
(text-fig. 223); 28 had twelve secondary tentacles, four inter-
vadial and eight adradial (text-fig. 224); and 3 possessed sub-
vadial tentacles in addition to these, the medusa illustrated in text-
fig, 225 (p. 1052) having as many as eight of these structures, two
in each quadrant, situated between the interradial and adradial
tentacles,

The sketches referred to above, although considerably enlarged,
are drawn to scale; they exhibit the fact that this variety of
Merisia lyonsi follows the general rule for multitentacular
meduse, in that the number of secondary tentacles increases
with the size and age of the individuals, The sequence of
the development of the secondary tentacles is also quite normal
in the specimens just described,

Although the majority of the multitentacular specimens of
Merisia exhibited a perfect radial symmetry and developed
their secondary tentacles in a perfectly normal order, a few
medusz proved abnormal in this respect and showed marked
asymmetry. One specimen had a single interradial tentacle
developed between two perradial ones, whilst another bore
interradial and adradial tentacles also in a single quadrant only.
Again, two meduse had these organs developed in two of the
quadrants (text-fig, 226), and a third had secondary tentacles in

* H.e. Stomotoca dinema Agassiz, and Orchistoma pileus Lesson : see Mayer (10)
pp. 111 and 219.
+ T'wo of these specimens were not included in the Table on p 1046.

MR, C. L. BOULENGER ON

1052

Text-fig. 225.

ary tentacles.

acular medusa with twenty second

Multitent

Asymmetrical medusa bearing secondary tentacles in two quadrants only.

VARIATION IN A JELLYFISH. 1053

three quadrants. This specimen had interradial, advadial and
subradial tentacles in two of these, whilst the third quadrant
carried only a single adradial one *.

Giinther has shown that in Limnocnida [9], tentacles of a
particular order are often fully formed in one quadrant before
there is any trace of them in the others. Browne also has called
attention to the fact that in Podocoryne carnea [7] the young
medusz do not always leave the hydroid colony with the same
number of tentacles, all have four perradial ones but the number
of interradial tentacles varies, some having two or three instead
of four, and one specimen he noticed to have a single one only.

I do not think that in the case of the asymmetrical medusz of
Merisia to which I have just referred, we are dealing with any
retardation in the development of the tentacles in certain
quadrants ; some of these individuals were of considerable size
and the gonadial diverticula well developed (ef. text-fig. 226), yet
certain quadrants showed no signs whatsoever of secondary
tentacles, although other quadrants possessed tentacles of the
third or even fourth order.

These five asymmetrical meduse clearly indicate that each
quadrant is capable of forming secondary tentacles quite
independently of the other quadrants of the bell. That this can.
occur in a radially symmetrical animal is distinctly stated by
Bateson [15] who remarks, that in radial series ‘as in Linear
Series, Variation, whether Meristic or Substantive, may take
place either in single segments (quadrants, sixths, etc.), or
simultaneously in all the segments of the body.” This statement
was based on observations made on two meduse, Clavatella
(Hleutheria) prolifera and Aurelia aurita.

Clavatella is a medusa which normally bears a single ocellus
at the base of each of its six tentacles. Claparéde [16] has called
attention to the fact that these ocelli are sometimes doubled ;
this duplicity may occur at the base of a single tentacle or
occasionally at the base of each tentacle instead of one.

Bateson also quotes the observations made by Romanes [4, 5]
on Aurelia aurita. In this form, in addition to changes symmetri-
cally carried out in the whole disc, one or more quadrants may
vary independently. Thus one specimen is figured in which two
quadrants are normal (7.e., each possesses one generative organ
and a set of radial canals) but the other half-disc is divided into
three. Similarly a particular quadrant may possess two sets of
organs or even three, the other three quadrants being normal or
nearly so.

In addition to the forms described above, three other medusze
showed abnormalities of a quite different type, which, although
known to occur in other genera, I consider worth recording.

In the first of these specimens, which was normal as regards

* This specimen is one of those not included in the Table published in my
original paper on Merisia lyonsi.

1054 MR. C. L. BOULENGER ON

the number of its tentacles, one of the latter organs was branched
in such a way as to present the appearance of a smaller tentacle

growing out laterally from a perradial one.

Text-fig. 227,

0:
oo

Diagrams showing the arrangement of tentacles in a normal and
five abnormal medusee of Merisia lyonsi.

P. Perradial tentacle. J. Interradial tentacle. S. Secondary tentacle in an
adradial position. ‘The subradial tentacles are not lettered.

VARIATION IN A JELLYFISH. 1055

Such branched or bifureated tentacles have been recorded pre-
viously by several authors, among whom Agassiz and Woodworth
[2| have described the phenomenon in medusz of Obelia (Hucope),
and Hargitt [8] came across a similarly abnormal specimen of
Gonionemus.

It seems highly probable that such abnormalities are not
congenital, these bifurcated tentacles having no doubt arisen as
the result of injury to normal ones.

Text-fig. 228,

Abnormal medusa in which the apicai canal is retained.
}

The abnormality presented by the other two medusz is of
greater systematic interest: each of these specimens possesses a
well-developed apical canal which projects from the base of the
stomach into the jelly at the summit of the umbrella (text-
fig. 228). This peculiarity is obviously due to the fact that these
specimens have retained the greater part of the canal which in
early life connected the cavity of the medusa-bud with that of the
parent-hydroid.

The presence of an apical canal was at one time considered to
he a specific character of some importance, but recent systematists *
have shown that such a canal occurs frequently as an individual
variation in many species which normally lose this organ in the
adult stage.

List of References.t

1. Boutencer, C. L.—“ On Meorisia lyonsi, anew Hydromedusan
from Lake Qurun.” Quart. Journ. Mier. Sci., vol. li. 1908,
pp. 357-378.

* (Cf, Mayer's remarks on Sarsia prolifera, and other species of this genus (10).

+ his list includes only those memoirs actually referred to in the text of this
paper; for a more complete bibliography I must refer the rcader to Dr, Mayer’s
recently published monograph (10).

1056 ON VARIATION IN A JELLYFISH.

2. Acassiz, A., & Woopworru, W. M.—‘“Studies from the
Newport Marine Laboratory: XL. Some Variations in the
Genus Hucope.” Bull. Mus. Comp. Zool. Harvard Coll.,
vol, xxx. 1896-7, pp. 121-150.

3. Mayer, A. G.—‘ The Variations of a newly-arisen Species of
Medusa.” Mus. Brooklyn Inst. Sci. Bull., vol. i. 1901,
pp. 3-27.

Romangs, G. J.—‘‘ Varietiesand Monstrous Forms of Medusee.”
Journ. Linn. Soc., Zool. vol. xii. p. 527.

Romangs, G. J.—Loc. cit. vol. xiii. p. 190.

Agassiz, A.— Meristic Variation in Sarsia.” Mem. Amer.
Ac. Sci. vol. iv.

Browne, E. T.—‘“‘ On British Hydroids and Meduse.” Proc.
Zool. Soe. London, 1896, p. 459.

. Harerrr, C. W.—‘* Variation among Hydromeduse.” Biol.

Bull. Wood’s Holl. Mar. Lab., vol. 11, 1901, pp. 221-255.

. GintHER, R. T.—* Zoological Results of the Third Tan-
ganyika Expedition conducted by Dr. W. A. Cunnington,
1904-1905.— Report on Limnocinida tanganice ; with a Note
on the Subspecies from the Victoria Nyanza.” Proc. Zool.
Soe. London, 1907, p. 643.

10. Maver, A. G.—‘‘ Medusz of the World.” Carnegie Inst. of

Washington, Publ. No. 109, 1910, vols. 1.-111.

11. Browne, E. T.—‘ On the Variation of the Tentaculocysts of
Aurelia aurita.” Quart. Journ. Micr. Sci., vol. xxxvii.
1894-5, p. 245.

12. Browne, KE. T.—“ Variation in Auvwrelia.” Biometrika,
London, vol. i. 1901.

13. Gorrz, A.—‘ Vergleichende Entwicklungsgeschichte der
Geschlechtsindividuen der Hydropolypen.” Zeit. f. wiss.
Zool., vol. Ixxxvii. 1907.

14. Bounencer, C. L.—‘ On the Origin and Migration of the
Stinging-Cells in Craspedote Meduse.” Quart. Journ. Micr.
Ste, VO lhe UGIO, yo, (ike

15. Batrreson, W.—Materials for the Study of Variation. 1894,
pp. 422-429.

16. CLiararkpE.—Beobachtungen iiber Anatomie und Entwick-
lungsgeschichte Wirbelloser Thiere, 1863, p. 5.

Oo OM OO

EXPLANATION OF PLATE LIX.

Fig. 1. A transverse section through a normal adult medusa of Merisia lyonsi.
The section shows the four radial canals and gonadial diverticula, as well as
the four endodermal ridges in the manubrium.

end., endodermal ridge. g.p., gonadial diverticulum. /am., endodermal
lamella. man., manubrium. ~7.c., radial canal.

Fig. 2. A similar section through a pentamerous specimen of the same species.
Note that although five radial canals and gonadial diverticula are well
developed, only four endodermal ridges occur in the manubrium,
Cf, p. 1048.

Lettering as in fig. 1.

12) Ay Sy USL IPE AS

Huth, Lith? London.
MARGINAL PROCESSES IN LAMELLIBRANCH SHELLS.

ON MARGINAL PROCESSES IN SHELLS. 1057

48. The Marginal Processes of Lamellibranch Shells.
By Cyrim Crosstanp, F.Z.8.

[Received May 27,1911: Read June 27, 1911.]
(Plate LX.* and Text-figs. 229 & 230.)

The existence of the ornamental projections on the surface of
so many lamellibranch shells is rather puzzling to the student
of Bionomics. Especially to one interested in the rate of growth
of mother-of-pearl sheli does their formation seem a sad waste of
shell-making energy.

The drawings given show examples of these processes in young
and adult shells. Inspection reveals one striking fact, that the pro-
portion of shell-building energy thus expended is very much greater
in quite young shells than in the mature ones of several species.
Compare, for example, the figure of a specimen of Margaritifera

Text-fig. 229.

Shells of Murex ramosus and upper valve of adult Chama foliata.
Half natural size.

The illustration will serve for identification of these common shells by
non-conchological readers.

margaritifera (the large mother-of-pearl oyster) seven months old
and that of the portion of the margin of one 54 years old, also the
young of Chama foliata and Chama sp. of about the same age
with the adult shells, and it is seen that relatively the processes
are gigantic in young shells (Pl. LX. and text-fig. 229). They
must have their use, or they could not be so greatly developed at
this, the second, critical stage of the bivalve’s existence Tf (the first

.* For explanation of the Plate see p. 1061.
+ Utility is suggested also by the fact that such processes cannot be regarded as
ancestral features such as in some other cases are more prominent in the young of a
species.

1058 MR. CYRIL CROSSLAND ON

critical stage is that during which the free-swimming larva must
attach itself to a suitable substratum or perish). At this second
stage the Aviculide at least are in the most active state they
attain to, and most other normally sessile lamellibranchs have
well-developed and actively used feet. Zridacna gigas is very
active at this stage, to mention a form probably not often seen
when so young, and particularly well fixed down when adult.
Their future life depends upon their success in crawling into the
position adapted to the special needs of their species. Also they
are now large enough to be attractive as food to fishes and erabs,
but not strong enough to resist claws and pincers. In the case
of M. margaritifera, among other species, the animal has little
to fear after it is one year old, being then too strong-shelled for
ordinary fishes and crabs.

Text-fig. 230.

Two specimens of Avicula zebra attached to Millepora alcicornis, and in the centre
of the latter a specimen of another genus of the Aviculidee which is more
common on corals. Another species of Avicwda is found on “black coral,”
Antipatharia. Half natural size.

Balistes flavimarginatus and B. viridescens, the largest of a
genus of shell-eating fish, are fairly frequently found here * in the
Red Sea, and occasionally break up ‘“ pearl” oysters two or three
years old, but those dissected were found to have preferred smaller
and weaker-shelled species. The big Rays are not common, and

* Dongonab, Port Sudan.

MARGINAL PROCESSES IN SHELLS. 1059

I have not certainly traced any damage to them. T'ridacna,
Spondylus, and Chama similarly have every chance of reaching
old age once the younger stages are passed, and the majority of
specimens found are thick and heavy ; on the other hand, Mar-
garitifera vulgaris (the Ceylon pearl-oyster), Ostrea sp.%, and
Avicula zebra of a year or two old are common, but above this
age are scarcely ever met with in the Red Sea. In the only
fishery for Margaritifera vulgaris which I have seen here, shells
two ox three years old, and small for that age, were being obtained
in comparatively insignificant numbers.

That energy so valuable elsewhere should be wasted upon un-
necessary appendages is also negatived by the consideration that
in this class, as elsewhere in nature, the struggle for existence is
severe, demanding the nicest adaptations to environment. This
is shown by the existence in this one bay of three distinct species
of Margaritifera, anatomically scarcely distinguishable, apparently
living together in the same habitat and obtaining food in
exactly the same way. Only after careful study does it begin to
be seen what delicate adaptations to special environments keep
the three species distinct, each in its own niche in the world.

Observations of the relations between living examples of the
following species and their common foe, Murex ramosus (text-
fig. 229), shows that these ornamental processes have a simple and
essential use to their possessor; indeed, without them they would
fali such easy victims to predatory Prosobranchs that those species
not otherwise protected must become extinct.

The following species living in exactly the same surroundings,
in the same artificial way, were observed, and their mortality
from attacks by Murex noted :—

Ostrea sp.% ...... No spinous processes. _ Decimated.
Avicula zebra... - oy 55
Margaritifera Processes small and Killed in large
mauritir. weak. numbers.
Margaritifera Processessmall,except Keptalive by frequent
vulgaris. in quite young shells. removal of Jurez,
otherwise numbers
perished.
Margaritifera Processes large and Attacked only excep-
margaritifera. strong, remain well tionally.

developed to at
least 6 years old.

We see that in these five species the liability to attack by
Murex (and probably by other prosobranchs, the behaviour of
which I have not observed) is just in proportion to the develop-
ment of spines. ‘The cases of attack upon J/. margaritifera
which I have seen corroborated, occurred only in stunted shells
in which these processes were ill-developed.

In the Aviculide at least the processes are made from the two

1060 MR. CYRIL CROSSLAND ON

outer shell-layers only, and as both of these are weak and horny,
so are the processes. These consequently get worn off the older
parts of the shell, and as increase in diameter becomes slow so
does the development of these processes* and in full-grown
shells practically ceases altogether.

It is obviously an advantage to the “oyster” to grow up
rapidly, and form a shell large enough to resist teeth and pincers
as quickly as possible. MMargaritifera vulgaris succeeds excellently
in this, attaining nearly its full size in a year; but it does this at
the expense of the sooner losing its shell-processes and so being
open to attack from J/urex, and doubtless other predatory
prosobranchs.

In MW. margaritifera also growth is extremely rapid during
the first year, and at the end of it the shell is strong enough
to resist such crabs and ordinary fishes as infest this bay; but
after the second year it is slow, a shell six years old being far
from full-sized. Consequently the formation of marginal pro-
cesses continues later, and with it partial immunity from attack
by durex. Sexual maturity is reached by all these species alike
in the second year.

From the result of my experiments recorded above, one wonders
how the first three species can possibly survive in Nature. The
explanation is very simple: they choose a habitat out of reach
of their foe. Ostrea sp.? and Margaritifera mauritiit inhabit
crannies among other shells £ or in stones, too narrow for Murex
to enter; while Avicula zebra, like other species of the same
genus and a few other genera of the same family, possesses a
foot that fears not the stings of corals, nor even those of J/ille-
pora, and so attains a habitat inaccessible to all others (text-
fig. 230, p. 1058).

Avicula zebra is very common on Millepora alcicornis in the
Red Sea; another advantage of this habitat is support above the
sea-bottom, clear of all obstructions to its respiratory circulation,
which latter is of greater value than would be supposed.

Given protection from enemies, and from being smothered in
mud, all these species will flourish wherever the experimenter
likes to place them, these two considerations being all that
causes the restriction of their habitats in nature.

* Only in a general sense is this true. In two specimens of M. margaritifera
of about the same age, viz. two years, one has added to its diameter much less
than has the other, owing to difference of conditions in which they grew ; but this
has not hindered development of processes, which are merely nearer.together in
the slower-growing specimens. It is when slow growth is due to age that the
formation of processes becomes imperfect.

+ This species, when over a year old, forms a postero-ventral non-nacreous addition
to the shell, similar to, but smaller than, that found in many other Aviculidz,
and which reaches extreme development in Mallews. In all, the addition is an
adaptation to enable the animal to inhabit narrow crevices, and its function is
similar to that of the siphons of those lamellibranchs which live buried in sand, or
in burrows.

{ H.g., on the under sides of large specimens of mother-of-pearl shell brought in
by the divers.

MARGINAL PROCESSES IN SHELLS. LO61

The way in which these processes protect the lamellibranch
has yet to be explained. From actual and repeated observation
one would expect that the prosobranch could extend its proboscis
for a short distance and insert it amongst the processes, which
are not generally very near together, and so drill its hole without
trouble. It is, however, necessary for the Murex to obtain a
very firm hold with its foot before it can operate its drill, and
one finds cases where a distinct impression of the foot has been
made among the growths on the shell before any damage has
been done at all. The point of special interest to the naturalist
as well as to the practical pearl-fisher, is that one cannot estimate
the damage done by Murex by counting how many of a lot of

dead shells have the characteristic hole drilled through them.
In move than three cases out of four no hole is drilled at all,
Murex having found an easier way. It finds the flexible edge of
the shell, then by contractions of its foot breaks a piece away.
The mucus of the foot-glands is then poured out in quantities,
and this has some poisonous effect, as the animal, while still
untouched, ceases to respond to the stimuli which ordinarily
cause a smart closure of the shell. The abundant flesh of this
species is all devoured before putrefaction sets in.

EXPLANATION OF PLATE LX.

Except for figs. 6 and 7, specimens of about the same age have been drawn to the
same scale, the age being something under six months, the magnification 3.

Fig. 1. Chama foliata. A dead shell, widely gaping, seen from above, i.e. from
anterior end. a, hinge; b, ventral edges of shell.

The shell-processes are seen to be about half the diameter of the shell in
length. At this age they and the shell ave brilliantly coloured with dark
red bands on a lght yellow ground. In the adult shell the processes
remain conspicuous (hence the specific name), but in the young they are
out of all proportion to the size of the animal that forms them.

Fig. 2. Upper valve only of Chama sp.? Shell red, processes white. In adult shells

processes quite inconspicuous, generally absent. | Animal then protected by

the extreme solidity of its shell.

Fig. 3. Spondylus sp.? A specimen which is beginning to take the adult shape

and to form the characteristic strong spatulate processes instead of the

brittle needle-lke serrated processes characteristic of the younger stages.

These serrulations are shown in oniy a few instances in the figure.

Fig. 4. A rather younger specimen of the same, with long slender processes only and
edge of shell serrated. Upper valve only drawn.

Fig. 5. Margaritifera margaritiferu. A specimen 12 mm. across shell, with

processes 7 mm. long. Probably three months old; younger specimens

than this have perfectly smooth shells like the umbonal region of the
present specimen.

Fig. 6. The same, X2, aged about 7 months. Showing development of processes
characteristic of the first two or three years’ growth. Some of the radial
rows of processes are thick and spatulate; others, which are thinner, shrivel
and become pointed, as in the figure, on drying.

Fig. 7. Thesame. Part of the edge of a three-year old shell which has grown slowly.

Shells more rapidly developed are distinguished by the wider spacing

of the processes.

I
bo

Proc. Zoou. Soc.—1911, No, LX XII,

1062 MR. CYRIL CROSSLAND ON

Warning Coloration in a Nudibranch Molluse and in
a Chameleon. By Cyrit Crossuanp, F.Z.8.

[Received June 12, 1911; Read June 27, 1911. ]

I. WARNING COLORATION IN CHROMODORIS.

Since the discovery of warning coloration there has been a
tendency toattribute protective value to displays of brilliant tints
which further observation has rendered doubtful, and the instances
in which actual protection has been experimentally proved are not
so numerous but that one more has some value.

The Chromodorids are, as their name implies, a family of Nudi-
branchs characterised by the development of colour shown by all
its members. Having collected a large number during the past
ten years, I may say that none of the family has been wanting in
this characteristic, except perhaps Casella atromarginata, which,
though handsomely, is not brilliantly, tinted. In this I merely
corroborate the general experience of marine collectors. Besides
this universality of colour the family is well defined structurally ;
indeed for the class the structure is remarkably uniform. The
usual depressed form of body, more or less ample margin, broad
foot, and feathery, usually tripinnate gills, are, in the more typical
Chromodorids, replaced by the opposite characters. A narrow foot
underlies a high body, the mantle is a mere shelf along the top
of its vertical sides, and the gills are simply pinnate *. Internally
the stomach is entirely embedded in the liver.

here is a smaller section of the family in which these char-
acters are less marked, the body having a wider mantle; in some
species a few of the gills are huaralived), and the Promveli Is partly
free from the liver, ‘aun indicating « connection with the ordinary
forms.

Otherwise structure is so uniform throughout the group, that
without detailed description of the colours of the living animal
identification of the species is impossible. Even with this deserip-
tion in full, determination of species is difficult, variation being
very Conetiemible so that, e.g., what one observer sees as black
bands on a white ground is to another white lines on a_ black

body ft.

* Arve these simply-pinnate gills the primitive form of Nudibranch gill, a modifi-
cation of the prosobranch ctenidium The highly specialised Chromodovids supply
the answer, since it is the less specialised members of the family that have the more
complex gills. A detailed study of the Chromodorid gill would show that its struc-
ture is more complex than the term “ simply pinnate ~ leads one to suppose, and
possibly would show details of higher organisation than do the tripinnate feathery
eills of ordinary Dorids. In the dev elopment of these gills from an originally irregular
vascular flap of skin, the tripinnate arrangement would be the earlier stage to be
reached.
+ Chromodoris quadricolor Eliot, Journ. Linn. Soc., Zool. xxxi. Nov. 1908, p. 107
= 0. elizabethina, P. Z.S. 190 L, p. 392, Pl. xxiv. fig. 4. Some other Chromo-
dorids are here figured including Cn igrostriata, a variety of the C. diardii
referred to later in this paper and a good example of the variation now
discussed,

WARNING COLORATION, 1063

The family is therefore probably monophyletic (so far as any
family can be), and it would seem that the power of colour
production is a single character, which, in a few of the cases in
which it has arisen, was associated with distastefulness or other
protective quality, and so, possessing a great survival value, led to
the foundation of the family. What colour or pattern might be
produced selection cared not, so long as it were conspicuous in the
eyes of nudibranch enemies, and duly correlated with inedibility *.

Hence the multitude of patterns and colours found in this one
class, and the variation above mentioned. Possibly one colour
is much the same as another in the biochemistry of the animal,
though so different to the human eye T

The survival value of the devices is evident from the abundance
of species in tropical waters. In collections, while species are
numerous, Individuals are usually rare, but the peculiar habits of
many nudlibranchs, which suddenly appear in local swarms and
shortly disappear again completely, show that as in, e. g., the case
of the Sharks of St. Andrews Bay %, which were not known to
exist there until new fishing methods were introduced, the pro-
portions in which species occur in collections are not at all those
in which they live in the sea, Of the Chromodorids [ remember
par ticularly hs annulata, C. sykesi, C. hilaris. and Ceratosoma
corniyerum as occurring for a few days in quite local swarms,
then dis sappearing absolutely ; while during the past fortnight
a hundred specimens of C. reticulata | PI. axel (p. 1068)] could
quickly be collected within a few yards of where [ write, but
to-morrow there may be not one.

The collection would be the more easy in that this species like
all Chromodorids crawls about in the open instead of skulking
under stones or in crannies after the fashion of most of the class,
except of course at low tide when it would otherwise be killed by
the sun,

In contrast with this, the general inconspicuousness of most
Opisthobranchs culminates in many cases in adaptations which
result in resemblance to surroundings so striking and so well in
aecord with the observed habitat, that there can be no doubt as to
protection being afforded by them §,

* Amidst this diversity we find that in all Chromodorids the edge of the mantle is
specially coloured. In others the gills and rhinophores are coloured differently trom
the rest of the body, as in the case of C. diardii,

+ That the colour of marine animals is often connected with their essential physi-
ology or apparatus of reproduction is dicated by the many cases in which the eggs
of a species are deeply coloured with the principal body colours of the parents.
This is conspicuously the case with numerous species of Polychwta and Nudibranchs,
The Chromodorid hereafter described under the name of C. reticulata lays eges the
colour of the yellow bands on the mantle. The purple band secretes macus of the
same tint,so that probably the purple colour has its physiological use, as well as
warning colour value.

+ McIntosh, W.C., “ Notes from the Gatty Marine Laboratory,” Ann. Nat. Hist,
yol. x. p, 254, 1902.

§ I have never had the fortune to possessa properly stocked aqnarium in which
experiments on this side of the theory conld be undertaken,

e 7
fo

1064 MR. CYRIL CROSSLAND ON

Two interesting facts are worth recording of the coloration of
several species of Tectibranchs, which besides being close simu-
lacra of their environment in other respects, each occur in green
and brown varieties, simulating green or brown weeds, or at least
in one case, the old and young shoots of Zostera. Living speci-
mens of these species are not now within my reach or further
details might be given, and sketches. The same thing happens in
a Scy llaeid*, the The ager green and the brown forms of which are
found near together i in Chuaka Bay, Zanzibar, whilst greenish to
deep brown varieties occur here (Dongonab) in the Red Sear The
other point about the coloration of these forms (Scyllaeid and
Tectibranchs) is the occurrence in so many species of small brilliant
blue spots, sometimes ringed with yellow, the only conspicuous
thing about otherwise scarcely visible creatures. These may be
a recognition marks ” but are more likely to be glands &.

We have then an excellent case for the theory that the brilliant
colour of these animals has a protective value. They walk abroad
where others must creep and hide.

Distastefulness was proved by trying to feed fish with specimens
(mostly CO. reticulata, twice with CO. diardii) thrown from the
windows of my homsebanti The kitchen refuse, and the shelter
afforded from the sun had made the surrounding water populous
with fish, and on a calm morning every detail of their movements
was clear. The water was shallow, the bottom sandy, with a
covering of short ‘sea grass,” the usual habitat of several fish and
often of considerable numbers of Chromodoris reticulata, more
rarely of CO. diardii.

The Garfish, Belone sp., is the only one of these animals with a
specialised diet. It is ever on the watch floating near the surface,
herding together the shoals of “ sardines” (Zngraulis ? boelama),

* Crosslandia viritis Eliot, P. Z.S. 1902, pp. 64-68; C. fusca Khot, Journ. Lim.
Soc., Zool. xxxi. 1908, p. 90.

+ The green Zanzibar specimen was found on Zostera, which is less abundant in
the Red Sea and from which I have not yet obtained Als species.

t Other brilliantly coloured Nudibranch families are the Polyceride and Aeolide.
OF the former I have seen too few specimens to generalise, but one of the most beau-
tiful, the wine-red Plocamopherus ocellatus, was merely a lighter tint of the colour
of its environment, a deep red polyzoan on which it ted. There is some evidence that
it owes part of its ‘colour to its foodstuff! Another, Treveleyana crocea 2, isa little
slug-shaped beast of a brilliant yellow colour. It periodically occurred in great
swarms in Chuaka Bay, Zanzibar, in the open, not hiding in any way. The Aeolids
are all bright coloured, I believe, and all, or nearly all, gain protection by living
amongst hydroid zoophytes, and even can utilise the stinging- thread cells of the
latter in their own bodies.

Tritonids often live among zoophytes, in Zanzibar often among fleshy Alcyonaria,
and are fairly conspicuously coloured. But Melibe fimbriata, a large animal, in colour
and in the shape of its cerata and the processes it bears, looks extremely likea
fragment of drifting fucus-weed.

1 Biot, Sir C., “Marine Biology of the Sudanese Red Sea,” Journ. Linn. Soe.
xxxi. Nov. 1908, p. 104.

2 Bliot, Sir C., “ Nudibranchs from Hast Africa and Zanzibar,” P. Z. 5.1904, vol. ii.
p. 87.

WARNING COLORATION. 1065

occasionally making dashes into the shallow water and carrying off
stragglers. Yeteven elone dashes up for any bait thrown into the
water, though the portions that sink below the surface are left to
the numerous bottom feeders. Of these we have Chrysophrys bifas-
ciata, Pimelepturus cinerescens, and two other species which I have
not identified, and they will eat, or have become accustomed to
eating, almost anything, including shelled specimens of Margariti-
fera vulgaris which had been kept in formalin for three weeks, and
the guts of Balistes flavimarginatus and B. viridescens which had
been in the same preservative overnight. Pimelepturus eats the
fresh viscera of its own species, or the pure white cooked flesh of
Caranx sp. equally readily. But none of these five fishes will eat
either of these two species of Chromodoris; attracted by the splash
they dashed up to them as to any other bait, but one fish after
another, of each species, as the nudibranch sank, at once turned
away after just touching it. Examples of Chrysophrys took the
animals into their mouths, but at once dropped them undamaged.

The behaviour of Pimelepturus was particularly interesting. If
a specimen of C. reticulata, which is white underneath (but for a
few purple spots under the edge of the mantle), fell on the sand
wrong side up, the fish would at once attempt to take it. If, how-
ever, the slug fell right side up, so displaying its gorgeous mantle,
Pimelepturus, swimming over it a few inches away, took no more
notice of it than of a stone.

What actually prevented these numerous fishes, of five distinct
species, from swallowing the Chromodorids? One can hardly
believe that a living slug could have a flavour more powerful
than that of formalin! Moreover, these two species have no odour
sensible to human organs, as have so many marine organisms,
including the Chromodorid Ceratosoma cornigerwm, and the
Tritonid Melibe fmbriata. The behaviour of these fishes, which
had for some weeks been feeding upon all kinds of kitchen
refuse, and especially the fact that when presenting an wn-usual
appearance Chromodoris momentarily attracted them, disposes
of the objection that they merely refused to eat an object which
was strange to them. Indeed, the natural circumstances of the
case make it impossible that C. reticulata should be unfamiliar
to any of these fishes except Belone.

But that this objection is a weighty one andin all supposititious
cases of protective coloration must be carefully considered, the
following examples will show.

The camels of the southern part of the Red Sea Province get
desert grazing all the year round, and are not fed upon “ dira ”
corn (Sorghum) in the summer. The consequence is that a
southern camel has actually to be taught with much patience (for
he deserves all that Kipling said about him) to eat corn. The
feast may be spread before him, but eat it he will not, until his
owner, seizing him by the nostrils, pours the corn into his mouth
with the other hand, spite of his groaning protests. Indeed some

1066 MR. CYRIL CROSSLAND ON

camels never get beyond this stage, and however hungry hold their
heads in the air above the pile of corn, crying like spoiled children
until the owner comes to feed them.

Contrast with this the behaviour of my two tame gazelles. From
six weeks old to maturity they fed on milk, corn, and occasionally
native leguminous plants, When fully adult they were introduced
to bread, sweet biscuits, sugar, green corn-stalks, and the leaves of
Acacia tortilis, all of which they ate readily without the stimulus
of hunger. I once tried to feed a sick camel on stale bread, by
way of invalid diet, It was impossible.

Another case is rather like that of the pigeons that starved
rather than eat a strange seed. I tried to change the feeding
place of my fowls by throwing their corn a yard or two to one side
of the usual place in the direction of the new one. When called the
fowls rushed up, stopped dead at the accustomed spot, and nothing
would make them move a foot beyond it. We can pair the fowls
with the opposite case, as we did the camels with the gazelles.
The land crab Ocypode makes its burrows a little above high-water
mark and never more than a yard or two higher up. But where
the camels and fowls were regularly fed, the ground is full of the
holes of these crabs ten or twenty yards above their usual habitat.
Also, instead of confining themselves to their accustomed diet of
dead fish, these lively crabs are ready for any experiment, from
sweet biscuits to handkerchiefs, and some must now be living on
camel dung and corn.

There are no oysters in Jerusalem, and consequently a native of
that city cannot be persuaded to try one. He has no religious
prejudice, or other dislike, but “ No, I do not know them ” is his
sufficient reason.

IT. WARNING CoLORATION IN A CHAMELEON.

I havenot seen the fact recorded that the Chameleon (Chameleon
sp.) ean change its colour so quickly as to frightenadog. While
staying in Zanzibar my host’s fox terrier showed hostile interest in
a chameleon someone had brought into the house. The chameleon
invariably tried to run away when attacked, but those who know
the species can imagine the ludicrous ineffectiveness of a chame-
leon’s flight. In a few seconds the impossibility of escape seemed
to reach the animal's brain, when it at once turned round, opened
its great pink mouth in the face of the advancing foe, at the same
time rapidly changing colour, becoming almost black. ‘This ruse
succeeded every time, the dog turning off at once. Among the
natural leafy surroundings of a chameleon the startling effect of its
sudden change of colour would be much greater. Imagine a dog or
cat nosing about suspicious of the presence of a live animal, but
unable to see anything. Until almost touching him the chame-
leon sits close, secure in his mantle of invisibility. Then suddenly,

WARNING COLORATION, 1067

when the dog’s nose is within six inches of him, the sudden change
from invisibility to the conspicuous black body and great red
mouth! I think many human beings would be startled, and in
fact the natives of Zanzibar hold the chameleon in horror, and
cannot be persuaded that its bite is not deadly. I tried the effect
upon some of them on my way home. ‘The ordinary native will
show great caution, but no panic, when a chameleon is handled
by a white man. I presented my specimen wneapectedly to
several parties I met on the road, thus :——

On approaching, ‘‘ Good morning,” which was quietly responded
to.

Just after passing, “See what a nice creature is walking on my
back.” Shrieks, and a stampede of fifty yards.

The specimen showed the ordinary light and irritation reactions
which have been frequently recorded.

1068 SIR CHARLES ELIOT ON

50. Chromodorids from the Red Sea, collected and figured
by Mr. Cyril Crossland. By Sir CHarues Huiov,
JS oils Giese BIB gp de Zatse

[Received June 12, 1911: Read June 27, 1911.]

(Plate LXI.*)

The present paper is a pendant to the one preceding it, and
gives some account of three species of Chromodoris from the Red
Sea which, though not new, are worth notice as being either
varieties of known species or forms hitherto imperfectly described.
The matter which it contains is mainly due to Mr. Crossiand, and
I have contributed merely the identification of the species and
a few notes on their internal anatomy. ‘The first species, Chr.
reticulata, is the one on which Mr. Crossland made his interesting
observations regarding the warning coloration of the genus. My
own experience does not entirely support his statement that
Chromodorids do not hide under stones but show themselves in
the open. I have often found them (as well as Zrevelyana crocea)
under stones on tropieal beaches, where their vivid colours
harmonize wonderfully with the ascidians and sponges found in
the same lecality, so that the nudibranch which when isolated 1s
conspicuous, is almost invisible at home. But, as Mr. Crossland
points out, many species are known to arrive on the shore in
considerable bands for the spawning season, and perhaps all do so.
At this critical period, at all events, warning coloration must be
useful to them.

The following notes on three Red Sea species indicate that
there is considerable variation not only in colour but in the details
of the buccal parts. In what appears to be the same species, the
denticulation of the teeth and the shape of the elements in the
labial armature may vary, and thickenings of the rhachis,
amounting to rudimentary central teeth, may be present or
absent.

CHROMODORIS RETICULATA Pease, var. (PI. LXI. figs. 1-3.)

See Bergh, ‘‘ Neue Nacktschnecken ete.” No. iv., Jour. Mus.
Godeffroy, Heft xiv. p. 9 ff.; and Eliot, P.Z. 8. 1904, pp. 386~7.
My. Crossland’s notes on the living animal are as follows :—

“This species is of the soft flat kind. Rhinophores with rather
high cups. Gulls 9, hinder ones short, set in a simple circle open
iivsthiadl. Snbranchedl simply pinnate, motionless. Foot projects
slightly behind the mantle. Mantle-edge slightly or distinctly
wavy, according to the exteusion of the animal.

“ Colour. Greater part of the back a fine network of chocolate-
brown on a grey-white ground; laterally both network and

* Vor explanation of the Plate see p. 1072.

soe
A tayiay?

to}
Sh 0)

E.Wilson, Cambridge i

CisURVOIMUOIDKORUD SW IRCOUME AU Ise, TEU ID) PSylavAN,

CHROMODORIDS FROM THE RED SEA. 1069

ground- colour become yellow, forming an ill-defined yellow band
which passes into white. Round the mantle-edge are two deep
coloured bands, the outer orange-yellow, the inner deep violet.
There is a faint line of purple round the extreme edge of the
orange-yellow border. Foot and whole under side pure white
except that colours of mantle-edge are repeated on the under side.
The inner border of violet on upper surface is thickened in places,
and these points are those where the bending of the mantle-edge
is marked when the animal is half contracted.

“ Rhinophores brown with clear white lines along the per-
foliations ; gills grey with white limes on the edge of the folia
and specks elsewhere.

‘‘One specimen measured 5:0 em.x1°6 cm. when fully ex-
tended.”

Four specimens which have assumed a uniform dull plum-
colour, no markings being visible. In most other respects they
agree with Mr. Crossland’s description of the living animal, but
rlheTeS he gives the gills as only 9, I find that they vary from
12 to 18. It often happens that in Chromodorids with many
gills, the smaller plumes are not protruded from the pocket and
thus escape notice.

The labial armature consists of bent rods, but they are not bifid
as in other specimens which I have examined. In twospecimens
the formula of the radula was 63 x 50.0.50 and 65 x 68.0.68 in
the widest rows. The rhachis, as in the specimens described by
me from East Africa, bears triangular thickenings, in which the
base is not clearly defined. ‘The first laterals are as usual in the
genus, low, flattened, and bearing 4 denticles on either side. The
remaining laterals are rather stout, hamate, and bear 6—8 distinct
denticles. The-outermost laterals are lower and bear two or
three denticles on the apex only.

These specimens are referable to Chr. reticulata, but seem to be
a distinct variety characterized by (1) the shape of the rods in the
labial armature, which are not bifid at the tips; (2) the presence
of thickenings on the rhachis of the radula; (3) aviolet border to
the mantle. As will be seen from my previous descriptions of the
species, the coloration is variable.

CHROMODORIS TINCTCRIA Riippell & Leuckart. (Pl). LXI.
figs. 4 & 5.)

Rippell & Leuckart, Neue Wirbellose Thiere des Rothen
Meeres, p. 32.

A single specimen described by Mr. Crossland from the life as
follows :—

“ Chromodorid in shallow water on Zostera growing in sand.

“ Length $85 mm. Breadth of mantle 46 mm.

‘“‘ Body and foot are narrow, mantle more than ordinarily wide,
margin thrown into folds and very mobile.

“Whole animal soft and smooth-skinned, but there are soft
warts on the back, about § inch high and broad. Gills in a row

1070 SIR CHARLES ELIOT ON

shaped GO ona raised base, 18 in number, generally simple but
some are forked; one quite arborescent, bipinnate. Pinnules
rudimentary. The gills are completely retractile and gill-pocket
can close over them. Rhinophores fully extended; numerous
fine white lines om deep crimson ground, otherwise perfoliations
hard to see; completely retractile. Head distinct, with pro-
minent tentacles.

‘General colour greyish white, but this is plentifully sprinkled
with clear white opaque marks, so that general appearance 1s
white. Over the body this is covered wilh 4 a delicate network of
crimson, the mantle being sprinkled with clear spots of the same
colour. Edge of mantle bordered with a thin clear line of bright
orange- -yellow. A broad crimson line runs up each angle of each
gill rhachis.”

Mr. Crossland adds in a letter: ‘“‘ The margin is much more
ample than in other species known to me. I mars never seen one
before in which the undulations are so deep or keep in such
constant motion.”

Ritppell and Leuckart’s diagnosis is “Colore lacteo, pallii
margine sulfureo-limbato : dorso venis punctisque sanguineis
notato: branchiis 19 circiter, pinmatis.” They also say that the
branchie are ‘“ pyramiden-formig,” which corr esponds to Mr.
Crossland’s observation that the pinnules are rudimentary. The
coloration is sufficiently distinct to make the identity of this
animal with Riippell and Leuckart’s Doris tinctoria certain.

The preserved specimen is somewhat distorted but the breadth
of the mantle margin is still noticeable, and the shape is not that
of an ordinary Chromodoris. The internal characters appear to
be those usual in the genus. Though the outer surface of the
liver is purple wherever it is covered by the hermaphrodite gland,
yet the organ itself is of a deep black and leaves a str ong stain.

The labial armature consists of two dark purple plates.

The rhachis bears thickenings much as in Chr. reticulata, and
the radula is of a type common in the genus. The inner laterals
are low and flattened, the innermost bear 3-4 denticles on either
side; the rest are denticulate on the outer side only and the
number of denticles rises gradually from 5 (on the second and
third laterals) to 15 or more. The teeth at the same time become
tall and elegant in shape. There are about 60 laterals in all.
The marginals are denticulate on the tip only but are not
degraded to mere plates.

Curomoports tNorinata (?) Bergh. (Plate LAI. figs. 6 & 7.)

Bergh, Siboga, pp. 157-159.

A single specimen described by My. Crossland from the life
as follows :—

‘¢? CHROMODORIS sp),

Shallow water say 3 feet deep, sand and “sea grass,” in
Dongonab Harbour (March 1911),

CHROMODORIDS FROM THE RED SEA. Og

“ Full length, from edge of mantle in front to tip of foot behind
82 mm., breadth across mantle 40 mm.; these being measured
when the mollusc is crawling. Turned over on its back, the body
is only 20 mm. wide, mantle remaining much as before ; its margin
is not only thus ample, but also mobile. It has no permanent
undulations.

“Right gills, simply pinnate and unbranched, the posterior two
short, the rest long even in proportion to the body. They are not
held stiffly as in so many Chromodorids but wave about in the
water with its motion. (Iam not sure whether there is any active
motion.) Rhinophores straight and sharply pointed ; distal ones
two-thirds perfoliate. They and gills completely retractile.

“ Colour. Back appears ‘of a deep red-brown with numerous
orange spots, small yet conspicuous to the naked eye. As the
orange shows itself elsewhere, we probably really have an orange
ground covered with a thick netw ork of purple (as seen under a
Jens) with small round meshes. This network ends 3 or 4 mm.
from mantle-edge, the inner half of the remainder is therefore a
deep orange. he outer half is violet, light proximally but almost
black in a narrow band distally. There is a very fine line of
primrose-yellow along the extreme edge of the mantle. The
under side of the animal is pure sihite. only the violet ef the
mantle-edge reappears on the other side.

‘« Besides the meshes of the network the back is marked by a series
of clear cut spaces each defined by a darkened line of purple.
These spaces, 2 mm, or so in diameter, may be oval, kidney-shaped,
or circular. Those which are near the orange band of the mantle
are a clear yellowish white, but the rest are clouded with orange
centrally and some have an ill-defined purple spot in the centre.

* Gills and anal papilla are orange on their inner surfaces (those
facing the centre of the cup they form), the rest white with a
little orange.

‘“* Perfoliate parts of rhinophores brown-red passing into purple
at tips. A yellow-white line is conspicuous along the anterior
side, running vertically, and very fine lines of the same run
horizontally on the posterior halves of the perfoliations.”

Mr. Crossland adds that the colour pattern was so complicated
and varied so much in different lights, that he abandoned the effort
of painting it and only made a black and white sketch.

In a letter he alludes to “the clear bilobed area behind the
rhinophores,” beneath which the eyes can just be seen.

The preserved specimen has become of a uniform plum colour
and shows no markings, but otherwise is as described by Mr.
Crossland. The branchie and rhinophores are both very oe
and both completely protruded. The branchie are only 8, but a
the base of the hindmost is a small tubercle which is ee
the beginning of a growing plume.

As in the last species the liver is of a very deep black, only
partly concealed by the hermaphrodite gland, The labial arm-
ature forms a complete circle, olive in colour, and consisting of

1072 - ON CHROMODORIDS FROM THE RED SEA.

bent rods, not strictly bifid but sometimes bearing asmall tubercle
near the tip. The formula of the radula is 70 x 80.0.80 as a
maximum, but it is considerably narrower in the anterior part,
which is also markedly darker in colour. There are thickenings
on the rhachis as in Chr. reticulata but even more distinct. The
first laterals are of the usual flattish shape and bear about
4 denticles on either side. The other laterals are rather tall, and
erect. They bear 8-12 very faint denticles, visible only under a
high power. The outermost laterals are irregularly shaped plates
with a few denticles at the top.

In most respects this animal closely resembles Bergh’s Chr.
inopinata. ‘The remarkable coloration is almost identical (unless
the light area behind the rhinophores proves to bea characteristic

.feature found in other specimens), but there are two differences.
In Bergh’s specimen (1) the branchiz consisted of five tufts each
subdivided into 3, 5 or 8 plumes, (2) the teeth of the radula bore

—7 distinct denticles. The first difference is hardly of specific
value by itself. Typically, the gills of Chromodoris are simply
pinnate but in some species they show a tendency to subdivide.
Still individuals belonging to such species have often normal
pianate gills. The difference in the teeth is more important. If
the denticles as drawn by Bergh are not exaggerated, we must
either recognize two species (the present animal being charac-
terized by its teeth, sumtplas gills, and perhaps some persistent
differences in wolleention) or else admit that considerable variation
in the vadula are possible. I am inclined to adopt the latter
alternative. It will be observed that though the appearance of
the teeth in the two specimens is different, this difference depends
not on an alteration of shape but in the greater or less develop-
ment of denticles. Thickenings on the rhachis seem to be
indifferently present or absent in several species.

EXPLANATION OF PLATE LXI.
Fig. 1. Chromodoris reticulata from a living specimen.
. Chromodoris reticulata. Dorsal view of the mantle and part of body in the
living animal; enlarged.
Chromodoris reticulata. (a) One of the lateral teeth from the middle of a
half row; (6) thickening on the rhachis.
. Chromodoris tinctoria from a living specimen.
Chromodoris tinctoria. Teeth: (a) third lateral, (6) lateral from the middle
of a half row, (c) outer lateral, (d¢) thickening on the rhachis.
Chromodoris inopinata (2) from a living specimen.
Chromodoris inopinata, a lateral tooth.

toe

ous

aug

Zo OMe elie

B.. Broom, del.

FOSSIL REPTILES FROM SOUTH AFRICA.

hei

jeyAS USMY, Je abesQnul.

R.. Broom, del,

POSSI BEPTILE SS’ FROM SOUTH AMRIC:®

ON NEW PERMIAN REPTILES FROM SOUTH AFRICA. 1073

51. On some New South African Permian Reptiles *.
By R. Broom, D.Se., C.M.Z.8.
[Received May 23, 1911: Read June 27, 1911. |
(Plates LXII. & LXIIL+)

Of the following new fossil reptiles a number have been
discovered by the Rey. J. H. Whaits of Beaufort West, a most
enthusiastic and successful collector. The others have been found
by myself. Though none of the species described represent any
strikingly new types, the series forms an important addition to our
knowledge of the Permian Fauna.

Suborder DINOCEPHALTIA.

MoscicPs CAPENSIS, gen. et sp.n. (PI. LXIT. fig. 1.)

This important new genus was discovered by Mr. Whaits and
myself on the farm Spitzkop in the Moordenaar’s Karroo.
Remains of a number of skeletons were discovered scattered over
and imbedded in an alluvial deposit about a rood in extent.
Though the remains were for the most part fragmentary and
weathered, and so completely mixed up that it is practically im-
possible to pick out the bones belonging to any one individual,
this matters the less seeing that all the skeletons appear to belong
to one species. There is considerable difference in size of the
bones, but I think there is reason to believe that this may be
accounted for by assuming that the small herd which perished
together comprised males, females, and inimature animals. As
portions of thirteen thigh ‘bones have been discovered, there must
have been at least seven individuals.

Owing to the scattered condition of the remains it is impossible
to reconstruct the manus aud pes and to give the exact number
of the pre-sacrai and caudal vertebrze, but with these exceptions
every detail of the skeleton is known. I hope shortiy to give a
full description of the remains in a monograph on the Dinocephalia.
In the present paper I shali merely give a preliminary desciiption
of the skull.

The skull, which I figure and which may be regarded as the
type, is, I believe, that of a young female. The left side is much
weathered and the whole skull considerably crushed, especially in
the occipital region, but otherwise the skull may be regarded as
nearly perfect. The greatest length of the skull is $45 mm., and
from the occipital condyle to the front of the snout 290 mm.

This new animal resembles Delphinognathus conocephalus Seeley
so closely, that it was only after very deliberate consideration
that I decided to place it in a new genus. That it is specifically

* On p. 1079 Dr. Broom proposes two new generic names, viz., Arcfosuchus and
Arctognathus.—ED1ror.
+ For explanation of the Plates see p. 1082.

1074 DR. R. BROOM ON NEW PERMIAN

distinet is beyond question, and the marked difference in the
temporal region seems to me of sufficient importance to Justify the
formation of a new genus for its reception,

The nasal region makes with the line of the teeth a sharp angle
of about 45°, and the snout 1s moderately broad. The pre-
maxillaries are small and each carried three teeth.

The maxillary is moderately largeand flat and carried 12 teeth,
of which the first four are large and may be regarded as canines.
The septo-maxillary is a small bone whieh ‘errs the lower wall of
the nostril, and passes backwards a short distance, separating the
nasal from the maxilla,

The nasals are fairly large rounded bones which are separated
from each other at the lower part by the internasal processes of
the premaxillaries.

The orbit is large and round, protected above by a marked
thickening of the Pupraorbital border and behind by a strong
postorbital arch. The bones surrounding the orbit appear to he
very similar to those of Delphinoc grits and Tapinocephalus,
though the limits of the prefrontal have not been clearly made
out. The lachvymal foramen is large and opens on the face.
The jugal forms the lower border of ‘the orbit. It isa flat and
relatively slender bone which passes back to meet the squamosal
and the quadrato-jngal.

The quadrato-jugal isasmall bone which rests on the descending
process of the squamosal and on the quadrate. It does not as in
Delphinognathus unite with the jugal to enclose a foramen.

The quadrate is very similar to that of the Pelycosaurs. In
the type it is considerably crushed and displaced.

The squamosal is large. It hasa long slender ascending process
which forms part of the posterior wall of the temporal fossa and
meets the parietal. A descending process supports the quadrate,
and an anterior meets the jugal. ‘The upper part of the squamosal
vests on a flat bone which forms part of the oeciput, and which
I regard as the opisthotic from a comparison with the bone in
Marsupials and Cynodonts.

The frontals are large and broad and enormously thickened.

The parietals are relatively small, but like the frontals extremely
thick. They enclose, as in Delphinognathus and Tapinocephalus,
a large pineal foramen. The edges of the foramen are elevated,
but the pineal region stands out less prominently than in
Delphinognathus.

The occiput is moderately flat, and the condyle large and
rounded. The upper side of the eondyle has a deep “hellow
evoove for the medulla, and in the middle line immediately below
the medullary groove is a small deep pit which passes forward
into the basioeeipital for the notochord.

The palate is imperfectly known, but appears te be fairly
similar to that of 7'apinocephalus.

The lower jaw is like that of Delphinognathus but more fully

REPTILES FROM SOUTH AFRICA. 1075

known. The dentary forms about half of the jaw, but owing to
a slender backward process which rests on the surangular it forms
about ¢ of the upper border.

The ‘splenial is slender. The angular forms the larger part of
the outer side of the posterior part of the jaw, and the slender
surangular the upper third. The articular, which has two con-
cavities for the quadrate, is a powerful bone which in front fits
in between the angular and surangular on the inner side of the
jaw.

The teeth are imperfectly known, but appear to be very similar
to those of Delphinognathus and HKecasaurus. This type closely
resembles the tooth figured by Twelvetrees and Seeley as the
tooth of Deuterosawrus, and if these authors are right in regarding
that tooth as belonging to Veuterosaurus, there can be no doubt
that Deuterosaurus is a Dinocephalian closely allied to South
African forms.

Suborder ANOMODONTIA.

DIALURODON WHAITSI, gen. et sp.n. (Pl. LXITI. figs. 6 & 7.)

This beautiful little Endothiodont skull was discovered by the
Rev. J. H. Whaits on the Beaufort West Commonage. It
is fairly complete, but the matrix is so hard that no development
has beenattempted. Fortunately, the skull was discovered broken
into quite a number of scraps, and, but for this, it would have
been regarded as belonging to a small species of Dicynodon.
The lower jaw was, however, broken across obliquely and the
fracture revealed the presence of a series of molars. A second
very badly weathered specimen appears to me to belong to the
same species.

The total length of the skull is 83 mm. From the beak to the
front of the orbit is only 22 mm., and to the back of the orbit
41 mm.,so that the orbit is entirely in the front half of the skull.
The ereatest width of the skull at the back part is about 50 mm.
The parietal region measures 19 mm. across its narrowest part, and
the frontal region is only 13 mm. across. Owing to the forward
position of the orbits the facial part is relatively small and also
narrow. The tusks are typically Dicynodont, but owing to the
premaxillaries beimg very small, placed more forward “than in
Dicynodon.

The molars are arranged in a row like those of Hndothiodon
uniseries Owen, but a second replacing set is seen developing on
the inner side of the functional teeth. There are probably about
8 teeth in use in each jaw. ‘The crowns are not displayed in any
of the teeth of the type specimen, but in the second specimen a
large part of one crown is seen. It is long and slender and has
coarse serrations on the anterior and posterior borders. In this
the teeth agree with those of Hndothiodon, and differ from those
of Pristerodon and Opisthoctenodon.

1076 DR. R. BROOM ON NEW PERMIAN

TAOGNATHUS MEGALODON, gen. et sp.n. (Pl. UXIT. figs. 2-4.)

This new genus and species is founded on an imperfect and
much crushed snout found by myself on the farm Kuilspoort,
Beaufort West district. Though little more than the orbital
region 1S preserved, this specimen is manifestly very unlike any
type previously known.

The frontal region is flat and moderate'y broad, the interorbital
measurement being 16 mm. ‘The orbit measures about 20 mm.
in length.

The greater part of the dentaries is preserved but the anterior
part of the beak is missing. ‘The jaw differs from that of all
previously known Anomodonts in the much longer portion covered
by horn, and also in that the anterior part of the lower jaw
instead of being narrower than deep, is here about twice as broad
as deep.

The teeth are remarkable in that while there is a tusk it
was probably relatively short, and in the lower Jaw there are two
large teeth each about half the size of the tusk. Not im-
probably there were two molars in the maxillary and two in the
mandible.

The nearest ally of Taognathus at present known is Prodicynodon,
but the affinity is not at all close.

OUDENODON BOLORHINUS, sp.n. (PI. LXIIT. fig. 10.)

This new species of Oudenodon was discovered by myself at
Kuilspoort, Beaufort West district. Unfortunately, the type
consists of only the preorbital portion of the skull, and this is
somewhat crushed and weathered.

The most striking characteristics of the species are (1) the
extreme shortness of the snout which brings the front of the
palate nearly under the orbit, and (2) the thickening of the nasal
bones to form a rounded boss which overhangs the nostrils.

The orbit is large and measures about 35 mm. in diameter.
The borders are thickened and rounded.

The maxillary bone is short but powerful; the caniniform
process being very massive. The upper part of the bone forms
part of the thickened lower margin of the orbit and nearly
sepirates the jugal from the lachrymal.

The premaxillary bone is also short and strong, and was
probably as represented in the restoration.

The nasals are short but greatly thickened, forming a large
median boss the sides of which overhang the nostrils.

The frontal bones are short and comparatively narrow, the
interorbital region measuring about 36 mm.

The only species of Oudenodon which seems to come near the
present one is Oudenodon strigiceps Owen, but this latter differs,
apart from differences that may be due to crushing, in the much
ereater size of the nostril and in the much more backward position
of the caniniform process.

REPTILES FROM SOUTH AFRICA, 1077

Suborder THEROCGEPHALIA.

AALUROSAURUS WHAITSI, sp. n. (Pl. LXITI. fig. 8.)

This new species of lurosaurus was obtained by Mr. Whaits
at Beaufort West. lurosaurus felinus also occurs at Beaufort
West on the same horizon, but the difference between the species
is considerable apart from size.

The specimen consists of the greater part of the left dentary,
much of each maxilla, the left premaxilla, and numerous other
fragments of the skull, besides a large number of. fragments of
the postcranial skeleton. The skeleton is so imperfect that no long
bone is entire, and most are represented by articular ends.
A few imperfect vertebrae are present, and a number of dis-
articulated bones of the manus and pes.

The symphysial part of the jaw makes a less obtuse angle with
the lower border of the ramus than in Wlurosaurus felinus.

There are four lower incisors and a single large canine. In the
type only the deep part of the root of the first incisor is left, so
that an accurate measurement of the space oceupied by these teeth
is impossible. The last is situated very close to the canine. All
the incisors are subequal and rounded. The canine measures
10 mm. x 6°5 mm. at the base; the height is not shown. The
molars are small and degenerate. The exact number is uncertain ;
three remain in the jaw, but apparently two have been shed and
replaced by bony tissue. Probably the young animal had 5 molars
(possibly 6). There is a large diastema of 15 mm. between the
canine and what is probably the occupied position of the Ist
molar, and the five molars havea space of 13mm. From the front
of the jaw to the back of the last molar is a distance of 48 mm.

The upper incisors are badly preserved, but the roots are
preserved and occupy a space of 21 mm. ‘The upper canine is
large and but shghtly curved; its anterior border is smooth and
rounded ; the posterior border is serrated. The antero-posterior
measurement at the base is 11 mm.

It seems not improbable that specimen R855 a in the British
Museum may belong to this species.

AKLUROSAURUS TENUIROSTRIS, sp.n. (Pl. LXITTI. fig. 9.)

This species is founded on a snout collected by myself at
Kuilspoort. It resembles lurosawrus whaitsc very closely, but
differs in that, though it is probably a rather larger form, the
snout 1s more slender and the mandible feebler.

The five upper incisors measure 24 mm.as compared with 21 mm.
in 4, whaitsi, and the whole measurement from 7’ tom’ is 65 mm.
in 4. tenwirostris as compared with about 52 mm. in 4. whaitsi.

Some of the lower incisors are well preserved. The first is a
rounded pointed tooth with a feeble serrated ridge on its outer
and posterior side. On the third incisor there is only a very
slightly marked ridge, and there are no distinct serrations, though
it is possible that they have been worn off by friction against

Proc. Zoon, Soc,—1911, No. LX XIII. Ce

1078 DR. R. BROOM ON NEW PERMIAN

the upper teeth. The molars are better developed than in
A. whaitsi.

The arrangement of the bones round the nostril is similar to
that in Scylacosaurus and Aloposaurus.

IcTIDOGNATHUS PARVIDENS, gen. et sp.n. (Pl. LXIT. fig. 5, &
IPL, TUMILIO US stor TT)

This new genus is founded on a small snout found by me at
Kuilspoort. It is, with the exception of Scaloposaurus constrictus,
the smallest known Therocephalian.

From the snout to the orbit measures 46 mm., and the whole
skull was probably not more than 90 mm. in length.

Owing to the weathering of the fossil, which was found in a
sloot and was thus considerably water-worn, the front of the
snout is badly preserved and the teeth are very imperfect. There
appear to be six incisors, all of small size. There is a fair-sized
canine with a very minute Ist canine in front of it. The molars are

numerous, Hight are preserved and two are probably lost, possibly

three. The dental formula would thus appear to be 7. Se. 2 Gdn =

The dental formula of Scaloposawrus was recently given by me as

5 @ 3 9 A suas ;
a.-,¢.-, m.~, the reason for believing that there are three canines

being that the maxillary series begins with oneminute tooth followed
by two larger than any of the succeeding molars. If we regard the
third tooth as a molar, the dental formula becomes the same as in
Lctidognathus. But though the dental formule may be the same,
there is no question but that the species must be placed in separate
genera, the large canine in the present specimen suflicing to remove
it from Scaloposaurus. ‘The canine measures antero-posteriorly
3mm.

ERIPHOSTOMA MICRODON, gen. et sp. n. (PI. LXIII. fig. 12.)

This small imperfect skull was found by Mr. Whaits at
Fraserburg Road. It is in two portions—a rather badly weathered
snout, and an equally badly weathered occipital portion. Though
the contact is missing, the two fragments can be united with fair
accuracy, and the whole skull as restored measures 110 mm. in
length. The snout is very flat and deep, measuring in the
canine region 27 mm. in greatest width, though the snout with
the lower jaw here measures 55 mm. in depth.

The incisors are long, narrow pointed teeth which are situated
well to the front, and are apparently only 4in number. Following
the last incisor is a long diastema of 13 mm. corresponding to the
position of the large lower canine. The canine measures 4°5 mm.
in antero-posterior length. The number of molars is uncertain.
Two only are preserved, and these are long, pointed, slender teeth.
The number of molars must, however, have been few.

The only genera to which Hriphostoma is nearly related are
Ictidosaurus and Lycosaurus. Ictidosaurus angusticeps, the only
known species, was described by me in 1903 from a specimen in the
South-African Museum. JLycosaurus was founded by Owen on

REPTILES FROM SOUTH AF'RICA. 1079

a badly weathered skull in the British Museum, which was made
the type of LZ. pardalis. Unfortunately, the genus Lycosaurus is
at present in considerable confusion owing partly to the bad
condition of the type, and partly to the fact that two other species,
which are not nearly related have been added to the genus.

In Lycosaurus pardalis, Owen determined the dental formula

BSS de ee : sins
to be t.3, ¢.;,™.z. Lydekker not only believes Z. tigrinus Ow.to be
correctly referred to Lycosaurus, but thinks itisreally indistinguish-
able from Z. pardalis. In my opinion the two do not even belong

to the same family. The dental formula of Lycosaurus pardalis
0 Pa 4 9 ° :
I determine as 7.-, c.-, m.. The minute canine in front of the

Jarge one is a character very frequently met with in the early
Therocephalians of the Pareiasaurus zone. The deep square
symphysis with the incisor teeth carried well to the front, and
the small size of the premaxillary portion are all early characters.
Owen gives the locality of the type as “‘Sneewberg mountain-
range,” but this is evidently a mistake, and Lydekker in his
Catalogue merely states that the specimen is from the ‘‘ Karoo
System of the Cape Colony.” It. is highly probable that the
specimen came from the Gouph.

Lycosaurus tigrinus is a much later type of Therocephalian.
Its dental formula is i, O.7, mo As it belongs to a different
genus, I would suggest the new generic name dArctosuchus to
contain A. tigrinus Owen.

Lycosaurus curvimola belongs to a still higher type of Thero-

cephalian, which is not nearly allied either to Lycosawrus or to
He ee 5
Ailurosaurus. Its dental formula is 7.-, ¢.-, m.-. I would

suggest for it the new generic name Arctognathus to contain
A. curvimola Owen.

Removing then these other genera and reserving Lycosaurus
for the type-species, we get a form resembling in many characters
Eriphostoma, but differing in being larger by a half, in having 5
incisors as against 4,and 2 canines instead of 1 as in Hriphostoma.
Lctidosaurus agrees with Lycosaurus as regards the incisor and
canine formula, but differs in having a large number of molars
and a number of other cranial characters. ‘The dental formule
of the three genera are as follows :—

° . )
Wichidosauiuss ee ae oe ee OS
Lycosaurus : pe ia asion 7 SCI Ba

? : 2
Pircpostonigls Vik CSEES ay eG a pean eas

The Geological Horizons of the Beaufort West Specimens.

Some years ago I endeavoured to subdivide the Beaufort series
into paleontological zones. At that time it was only possible to
do so on very broad lines. The area is so extensive, and except

on the upper part of the series there are no lithological characters
(ae

1089 DR. R. BROOM ON NEW PERMIAN
to correspond to the paleontological. Further, as wide areas of
the Karroo are covered by wind-blown dust which forms what
may be regarded as a sort of loess deposit, and the exposed
portions, except in the case of escarpments, are usually isolated, it
becomes a matter of extreme difficulty to connect up the various
beds. The shales for a thickness of 3000 or 4000 feet present no
distinguishing characters, and at present we can do little more
than collect fossil specimens and note the localities.

I subdivided the Beaufort into six zones which may be grouped
as follows :—

(6. Cynognathus Beds.
| 5. Procolophon Beds.
Middle Beaufort 4. Lystrosaurus Beds.
3. Cistecephalus Beds.
Lower Beaufort < 2. Hndothiodon Beds.
1. Pareiasaurus Beds.

Upper Beaufort

The Lystroswurus zone probably corresponds to the Lower
Triassic ; the Cynognathus zone to the Upper Triassic. The
Pareiasaurus beds are probably Middle Permian ; the Hndothiodon
and Cistecephalus beds are probably Upper Permian. <All work
done since 1905 has gone to confirm the correctness of the con-
clusions then arrived at, but it has shown that we will some day
be able to subdivide the zones into a large number of subsidiary
ZONES.

The Pareiasaurus zone is the oldest one where fossils are
numerous. Pareiasawrus is its most characteristic form, but there
are a host of other known genera, mostly of 'Therocephalians.
Small Anomodonts—Dicynodon and Oudenodon—are also not
uncommon.

Above the Pareiasaurus zone we have deposits which measure
about 2000 feet, principally characterised by the abundance of
Dicynodon and Oudenodon. Of every six fossils obtained, five are
those of Dicynodon or Oudenodon. As, however, these genera
occur at all horizons of the Beaufort, it seems unwise to speak
of this as the Dicynodon zone. I have therefore called it the
Endothiodon zone, as Endothiodonts are met with throughout it
and are known at no other horizon. 'Therocephalians are not
common, but may be met with at any height. Curiously enough,
with the exception of Propappus and Sawrosternon, no genera are
known from the Hndothiodon zone which do not belong to the
Anomodontia or the Therocephalia.

Above the Hndothiodon zone is a zone probably not more than
1000 feet thick, which is characterised by the presence of
Cistecephalus, and the higher types of Therocephalians. ‘The zone
is at present the least known of any part of the Beaufort, mainly
because since Andrew Bain’s time no collector has had an oppor-
tunity of doing much with it.

At Beaufort West, one has an opportunity of studying to
advantage the Hndothiodon zone. The township is situated on

REPTILES FROM SOUTH AFRICA. 1081

an undulating plain which is geologically probably not more than
500 feet above the Pareiasaurus zone. The subzone is charac-
terised by the presence of Hndothiodon uniseries Ow.,—a form
that at present is not known from any other part of South Africa.
In the Beaufort West district also occurs Hndothiodon bathystoma
Ow., but whether the horizon of this is above or below that of
FE. wniseries is at present unknown.

Near Beaufort West is the escarpment of the Nieuweveld
which rises a few miles north of the township to a height of 3000
feet or more, and though it is extremely difficult collecting in the
steep slopes of shale, a number of forms of interest have been
collected at various horizons.

As Dicynodon and Oudenodon occur at all levels, it might be
thought well to subdivide the zones by the species of Dicynodon ;
but there is a serious difficulty. Dicynodon is the most trouble-
some genus we have to deal with. Specimens differ so greatly in
size and shape that one hardly knows what to do unless one does
as was practically done by Owen, make every specimen the type
of a distinct species. For many years to come the genus
Dicynodon must remain in utter confusion, and will be useless for
stratigraphical work. On the other hand, the Therocephalians
will be as reliable guides as Ammonites and Trilobites are in the
marine rocks of Europe.

In the following diagram are represented the horizons of the
Beaufort West types described in this paper, and of others whose
horizons are known. It must, however, be understood that the
heights in feet are only approximate.

|

|

| 2000 feet ..........55....... Ictidognathus parvidens Br.: Allurosaurus tenui-
rostris Br.; Oudenodon bolorhinus Br.; Cistecephalus
microrhinus Ow.; Dieynodon sp.

| 1500 feet ..................... Aloposaurus gracilis Br.
| Dicynodon sp.

| 1000 feet ..................... DPaognathus megalodon Br.; Oudenodon sp.;
| Dicynodon sp.
500 feet ...............-..... Dicynodon sp.
Dicynodon sp.

Beaufort West horizon ... Hndothiodon uniseries Ow.; Dielurodon whaitsi Br. ;
Alurosaurus whaitsi Br.; Scymnosaurus sp. ;
Dicynodon sp.; Oudenodon sp. lurosaurus

felinus Ow.

It may be well to regard the Hndothiodon zone as reaching
1600 feet above Beaufort West, and then passing into the
Cistecephalus zone.

1082 MR. E. G. BOULENGER ON A

EXPLANATION OF THE PLATES.
Puate LXII.

Fig. 1. Side view of skull of Moschops capensis Broom. 2nat.size. Ang. Angular;
Art. Articular; Ju. Jugal; O.O. Opisthotic; Qu. Quadrate; Q.J. Quadrato-
jugal; S.Ang. Surangular: Sma. Septomaxillary ; Sq. Squamosal.

2. Front part of skull of Taognathus megalodon Broom. Nat. size.

3. Under view of lower jaw of Taognathus megalodon Broom. Nat. size. The
canine of the right side is seen in oblique section.

4. Section across lower jaw of Taognathus megalodon Broom. Nat. size.

5. Upper view of snout of Letidognathus parvidens Broom, Nat. size.

Prate LXIII.

Fig..6. Side view of skull of Dielurodon whaitsi Broom. Nat. size.
7. Section across lower jaw of Dielurodon whaitsi Broom, at the crack
indicated in figure 6. Nat. size.

8. Upper view of mandible of _4lurosaurus whaitsi Broom. Nat. size.
9. Side view of snout of Alurosaurus tenuirostris Broom. 2 nat. size.

10. Side view of snout of Oudenodon bolorhinus Broom. 3 nat. size.

11. Side view of snout of Ietidognathus parvidens Broom. Nat. size.

12. Side view of snout of Hriphostoma microdon Broom. + nat. size.

52. On anew Tree-Frog from Trinidad, living in the Society’s
Gardens. By Hpwarp G. BouLEencer, Curator of
Reptiles to the Society *.

[Received September 20, 1911: Read October 24, 1911.]

(Plate LXIV.+)

In July last the Zoological Society received from Dr. Lewis
H. Gough an interesting collection of Batrachians and Reptiles
brought back by ‘him from Trinidad. Among these I found
examples of three frogs which had not been previously recorded
from Trinidad, viz.: Hyla venulosa Daud., Hyla rubra Daud., and
one which is evidently undescribed, and for which I propose the
name of Hyla goughi, after its discoverer. This little Hyla, one
of the smallest of the genus, was fortunately represented by
numerous specimens, which have enabled me to observe the wide
and rapid changes of colour which this species undergoes, and of
which an idea can be gained from the annexed coloured plate
made by Mr. J. Green at the Gardens under my direction.

HYLa Goucut, sp.n. (Pl. LXIV.)

Tongue circular, slightly nicked and slightly free behind.
Vomerine teeth in two rounded groups between the choane.
Head slightly broader than long; snout rounded, a little shorter
than the eye, which is large and very prominent. Canthus
rostralis feebly marked. Loreal region very slightly concave.

* Communicated by the Secretary.
+ For explanation of the Vlate see p. 1083.

eZ) LONE wileplexgl Va

ad.Green del.et Chromo lith.

GOUGHS TREE FROG.
(HYLA GOUGHI.)

NEW TREE-FPROG FROM TRINIDAD. 1083

Interorbital region a little broader than the upper eyelid.
Tympanum fairly distinct, 4 diameter of eye, Fingers rather
short, with small discs, outer 4 webbed. Subarticular tubercles
feeble. No projecting rudiment of pollex. Toes 4 webbed.
The hind limb being carried forward along the body, ‘the tibio-
tarsal articulation reaches between the eye and the end of the
snout. Tibia half the length of the head and body. Upper parts
smooth, with minute granular warts on the head and the anterior
part of the back. Male with a feebly developed subgular vocal
sac, forming loose folds.

Coloration.—Vhe vapid changes in colour which this frog
undergoes are probably unparalleled in any other Batrachian.
The same individual may vary dorsally from dark brown, reddish
brown, various shades of yellow, to a very pale greyish white.
When startled the majority became of a bright lemon-yellow.
Jn one specimen I observed the head, fore limbs, and anterior
part of the body to be dark brown, whilst the posterior part of
the body and hind limbs were greyish white. In another speci-
men the right half of the body was brown, the left half greyish
white. A brown or grey marking, often hourglass-shaped,
edged with darker or lighter, extending from between the eyes to
the anterior third of the back, followed by one or two transverse
bars, is frequently present, appearing and disappearing with
great rapidity. In one specimen the warking took the shape of
a eross-bar between the eyes and two parallel longitudinal bands
extending along each side of the entire length of the back.
Faint cross-bars on the hind limbs are occasionally present.
Lower parts white or yellow. Iris golden, much obscured by
brown pigment; a clear golden line borders the pupil, which
when fully contracted becomes perfectly linear.

The largest specimen measures 22 mm. from snout to vent.

This little Tree-Frog is extremely agile in its movements,
making leaps of quite six feet. In the daytime it usualiy kept
quiet, sticking to the leaves in the terrarium. On arrival the
males issued a sharp creaky note, but became perfectly mute
after a day or two at the Gardens,

This species appears to be more nearly related to Hyla strigilata
Spix, from Brazil, and Hyla misera Werner (Zoo), Anz. 1903,
p- 252), from Caracas, Venezuela, than to any others. It differs
from both in the shorter snout ; ‘from the former also in having
more fully webbed toes; from the latter in having a smaller
tympanum, and a shorter web between the fingers.

EXPLANATION OF PLATE LXIV.
Gough’s Tree-Frog (Hyla goughi).

1084 MR. C. E. HELLMAYR ON THE

03. A Contribution to the Ornithology of Western Colombia.
By C. EH. Hetimayr, Curator, Division cf Birds,
Zoological Museum, Munich *.

[Received April 26, 1911; Read June 13, 1911.]

Page

Tc Introduction esas ce ee ee ee ee LOS:
If. Account of Mr. M. G. Palmev’s localities.................. 1085
Iif. Account of the Species .. SREY Ae tee dn AM, een 1086
Wik Coneluvsion sy peice We ree mn hed er ae meee ea toy EATON)

I. INTRODUCTION.

The following account is principally based on a collection of
birds made by Mr. Mervyn G. Palmer, one of Mr. W. F. H.
Rosenberg’s field collectors, in the second half of 1908 and in
1909. Though numbering hardly 700 specimens, the set is of
considerable interest, containing a large percentage of rare species
as well as several novelties which I have already deseribed, partly
in the ‘ Bulletin of the British Ornithologists’ Club,’ partly in the
‘ Revue Frangaise d’Ornithologie.’

The district of Western Colombia in which Mr. Palmer has been
werking had not been visited by many naturalists before, and,
putting aside some seattered notices in scientific periodicals, few
accounts have been published about the birds of this little-known
region. Our first knowledge is due to the exertions of the cele-
brated French traveller, Ad. Delattre, who, prior to 1846, made
considerable collections in the vicinity of Buenaventura (Chocé
Bay), in the Western Cordillera at Juntas, Cali, ete., in the neigh-
bourhood of Popayan, notably on the Puracé, as well as at Pasto,
near the Heuadorian frontier. The new species of Humming-
Birds discovered on this trip were made known by Delattre and
Bourcier in the ‘ Revue Zoologique,’ vol. ix., 1846, pp. 305-312,
while a number of birds belonging to other families are discussed by
M. de Lafresnaye in the same journal, vol. x., 1847, pp. 67-79.
The next important contribution is Cassin’s Catalogue of the Birds
collected by Lieut. N. Michler’s expedition to the Rivers Truando
and Atrato, just south of Darien, published in the ‘ Proceedings of
the Academy of Natural Sciences of Philadelphia’ for the year
1860, pp. 132-144, 188-197. This was followed by Sclater and
Salvin’s account of the late T. K. Salmon’s extensive collections
from the State of Antioquia, in the P.Z.S. 1879, pp. 486-550,
wherein 468 species are dealt with. In 1894, Mr. W. F. H. Rosen-
berg visited the Rio Dagua, working chiefly at Juntas and Cali, in
the Western Cordillera. His birds went to the late Adolphe
Boucard, who published a list of the Humming-Birds in ‘The

* Communicated by Dr. P. L. Scuater, F.R.S., F.Z.S.

BIRLS OF WESTERN COLOMBIA. 1085

Humming Bird,’ vol. v., 1895, pp. 5-7, while the bulk of the col-
lection was never reported upon. Mr. Gustav Hopke, im 1896 and
1897, sent a fair series from the same district to Count Berlepsch,
who described several new species in the ‘ Ornithologische Monats-
berichte,’ vol. v., 1897, pp. 173-176, and in the ‘ Ornis’ xiv., Feb.
1907, pp. 347, 361, 365. Mr. Eugene André, in 1899, forwarded
a large collection of birds, from the environs of Buenaventura
and the western slope of the Andes above that town, to Comte
de Dalmas, of Paris. Unfortunately, the greater part of 1t was
subsequently destroyed by accident, and merely a list of the
Trochilide published by Messrs. Simon and de Dalmas in ‘ Ornis,’
x. 1901, pp. 216-224. Lastly, Mr. Outram Bangs, the well-
known ornithologist of Boston, who had obtained two of Palmer's
earlier consignments, gave short accounts of the more interesting
species in the ‘ Proceedings of the Biological Society of Washing-
ton,’ vol. xxi., 1908, pp. 157-162,* and xxiii., 1910, pp. 71-76,
describing several supposed novelties some of which are undoub-
tedly distinct, while others prove to be identical with species
previously named.

The material discussed in this communication is deposited in the
collection of the Zoological Museum at Munich with the exception
of the few species given in brackets.

II. Account oF Mr. PAatmer’s LocALirtizs.

The whole of Mr. Palmer’s collection in possession of the
Munich Museum was secured in the province of Chocé, though
in two different districts. The bulk is from the hot, tropical
valley of the Rio San Juan and its tributaries, the Tamana,
Sipi, Condoto, Calima, Cajon, and Garrapatas Rivers, while a
smaller number of specimens was collected on the Pacific slope of
the Western Cordillera near the head-waters of the San Juan, at
altitudes of from 2800 to 8000 feet.

IT am indebted to Mr. Rosenberg for information about the
nature and altitude of some of the localities which he had visited
himself in 1894, They may be separated into three divisions :—

(A) Rio Dagua District.
Sad Joaquim, Bahia del Chocé (= Buenaventura of Huropean
maps).
El Paillon, several hours’ journey up the Dagua (HK. André).
San José, 600 feet (visited by André and Hopke).
Los Mangos (=Juntas, Rio Dagua), cirea 1000 ft. Hot, tropical
country.

* The statement that the material came from N.W. Colombia, “just south of
Darien,” is a mistake. I am informed by Mr. Rosenberg that the San Antonio
referred to isa village on the road from Buenaventura to Cali, just over the pass
of the Western Cordillera. M1. Bangs appears to have identified it with the San
Antonio, on the Rio Sucio, a tributary of the Atrato, where, however, Mr. M. G.
Palmer never was.

1086 MR. C. FE. HELLMAYR ON THE

Jiménez, 1600 ft., a morning’s walk from Los Mangos, in a
deep ravine by the side of the pass between Los Mangos
and a place called Ventanas, on the road to San Antonio
and Cali. Forest country. Above Ventanas the country

_ becomes open. .

Naranjo (1900 ft.) and Plano de los Monos (2600 ft.). (Visited
by HK. Andre.)

Naranjito, 3900 ft.

Pavas, 4400 ft; San Luis, Bitaco Valley, 4400 ft.

La Maria, 4700 ft.

Palmar, near La Maria.

San Antonio, R. Cali, 5400 to 5800 ft. On road from Buena-
ventura to Cali, just over the Pass of the Western
Cordillera. Mr. Rosenberg passed through San Antonio
many times, but never collected there. The first lot that
went to Mr. Bangs was chiefly brought together at this
locality.

(B) Rio San Juan District (main stream and tributaries). Hot,

tropical country covered with dense forests.

R. San Juan: Noanama, alt. 100 ft.; Tad6, farther north,
230 ft.

R. Cajon, a small affluent.

R. Calima: Guineo (sea-level).

R. Sipi: Sipi, 150 ft.; Rio Garrapatas, 150 ft.

R. Condoto : Condoto, 150 ft.

R, Tamana: Noévita, 150 ft.; El Tigre, 320 ft.; Juntas, 400 ft.*

(C) Pacific slopes of the Western Cordillera, sources of the Rio

San Juan.

Pueblo Rico, San Juan slopes, 5200 ft.

Siaté, Rio Siaté, near Pueblo Rico, 5200 ft.

KR. Jamaraya, one of the headwaters of the R. San Juan: Loma
Hermosa, 4180 ft.; La Selva, 4600 ft.

Tatama Mountain. Mr. Palmer collected at various stations
(2794 to 8000 feet).

III. Account oF THE SPECIES.f

1. TURDUS TRISTIS DAGUA Berl.

[Merula tristis Swainson, Philos. Magaz. (n. ser.) i. p. 869
(1827.—Temascaltepec, Mexico). |

Lurdus dague Berlepsch, Ornith. Monatsber. v. p. 176 (1897.—
San José, Rio Dagua, 8.W. Colombia); Hartert, Nov. Zool. v.
1898, p. 478 (Cachavi, N.W. Ecuador).

No. 1957. g ad. San Joaquim, Bahia del Chocé, 1.viii.08.—
Wing 107; tail 84; bill 18 mm.

* Not to be confounded with Juntas on the R. Dagua.

+ [The absence of brackets round the name of an author to indicate that his
species has been transferred to another genus is not due to Mr. Hellmayr, but is in
accordance with the custom of the Zoological Society —Ep:ror. |

BIRDS OF WESTERN COLOMBIA. ' 1087

Nos. 2078, 2180. ¢ ¢ ad. Sipi, ll.ix., 12.x.08.—Wing 110,
HOS ss tale Ser 95m bull S anne

No. 2299. g ad. Noévita, Rio Tamana, 16.xi.08.— Wing 106;
tail 78; bill 18 mm.

No. 2370. 9 ad. Ndévita, Rio Tamana, 5.xi1i.08—Wing 105;
tail 76; bill 18 mm.

No. 2072. 2 imm. Rio Sipi, 10.1x.08.—Wing 107; tail 81;
bill 17 mm.

“Tris, feet and bill dark brown.”

In spite of the late Dr. Sharpe’s doubts there can be no
question that 7’. t. dague is a perfectly distinct form, though, to
my mind, its relations are more correctly expressed by a trinomial
appellation. Compared with a good series of 7’. tristis cnephosa
Bangs, from Chiriqui and Western Costa Rica (Miravelles), the
Colombian birds have shorter wings and tail, a much shorter,
slenderer and darker (blackish-brown) bill, and differ also in
coloration. The upper parts are deep reddish sepia-brown (instead
of greyish olive or olive-brown), the breast and sides dark
rufescent brown (instead of pale brownish grey), and the sides of
the head deep sepia-brown (instead of sooty black).

Tt. dague is peculiar to Western Colombia and N.W. Ecuador,
whence Miketta and Flemming sent numerous specimens to
Mr. Rosenberg. Hcuadorian skins are practically identical with
the topotypes forwarded by Mr. Palmer. This Thrush is an
iwhabitant of the humid lowlands and does not occur in the
mountainous parts of the interior.

2. HyYLocICHLA USTULATA SWAINSONII Cab.

[Turdus ustulatus Nuttall, Manual Orn. U.S. & Canada, Land
Birds, ed. 2, pp. vi, 830 (1840.—Columbia River). |

Turdus swainsoni Cabanis in Tschudi’s Fauna Peruan., Aves,
p- 188 (1844-46. — “ New-Jersey, im Monat October,” coll.
Cabanis).

No. 2302. Q ad. Novita, 17.xi.08. “Iris dark brown, feet
pink, maxilla black, mandible light brown, tip black.”

Identical with other specimens from Bogota, Ecuador, and
Venezuela (Merida).

3. MYADESTES RALLOIDES D’Orb.

Muscipeta ralloides D'Orbigny, Voyage dans |’Amér. mérid.,
Oiseaux, p. 322 (between 1838 & 1847.—Chulamani, Yungas of
Bolivia).

Myiadestes ralloides Sclater & Salvin, P. Z. 8. 1879, p. 492
(Retiro, Concordia, Medellin, 8. Elena).

Nos. 3760, 3771. $ Q ad. Siaté, 5200 ft.: Sept. 17, 1909.—
Wing 84, 82; tail 75, 72; bill 1] mm.

“Tris dark brown, feet light brown, maxilla black, mandible
light brown.”

The specimens agree perfectly with others from Bogota, while

1088 MR. C. E. HELLMAYR ON THE

skins from Cumbre de Valencia, Venezuela, have decidedly smaller
bills. Typical Bolivian birds are not available for comparison.
M. ralloides is probably divisible into several geographical races.

4. HELEODYTES ALBOBRUNNEUS HARTERTI Berl.

[Heleodytes albo-brunneus Lawrence, Ibis, iv. p. 10 (1862.—
“On the line of the Panama Railroad near the summit of the
Atlantic Slope ”.)|

Heleodytes harterti Bevlepsch, Ornis, xiv. p. 347 (Feb. 1907.—
San José, Rio Dagua, W. Colombia).

No. 2522. g ad. El Tigre, Rio Tamana, 320 ft., 9.11.09.— Wing
92: tail 88; bill 23 mm.

No. 2523. @ ad. Same locality and date.—Wing 83; tail 78 ;
bill 20 mm.

“ Tris reddish brown, feet grey, bill black.”

The specimens agree in coloration with the types kindly lent by
Count Berlepsch, and have the back, wings, and tail very much
darker, more brownish black, than typical albobrunneus from
Panama. The dusky markings of the under tail-coverts are
less regular, more spot-like, and the bill is black instead of
pale brown. The difference in size, however, proves to be not
constant, the female from Chocé beimg scarcely larger than
Panama examples *. It may be mentioned that the female has
the head all round pure white hke the male, while that of albo-
brunneus is said to have the pileum clouded or streaked with
pale brown.

H. a. harterti is as yet known only from the Chocé district in
Western Colombia.

5. LEUCOLEPIS PH ZOCEPHALUS PH#OCEPHALUS Scl.

Cyphorinus (sie) pheocephalus Sclater, P. Z. 8. 1860, p. 291
(end of 1860.—Esmeraldas, N.W. Ecuador); Sclater & Salvin,
P. Z.S. 1879, p. 492 (Remedios, Antioquia).

Cyphorhinus brunnescens Sharpe, Cat. B. Brit. Mus. vi. p. 293
(1881.—“ Cauca Valley,” sc. Remedios).

No. 2555. Q ad. Juntas, R. Tamanda, 24.11.09.—Wing 66;
tail 32; bill 20 mm.

“Tris brown, feet dark brown, bill black.”

This specimen, a perfectly adult bird, agrees exactly with topo-
typical examples from N.W. Ecuador (8. Javier), and others from
Chimbo. Perhaps it is a trifle darker chestnut on the back and
slightly deeper rufous underneath. The type of C. brunnescens
Sharpe, which I have examined in the British Museum, is merely
a young bird of C. pheocephalus. Kcuadorian specimens in cor-
responding plumage are exactly like it. Moreover, it 1s a well-
known fact that in the species of this genus the young birds have
the upper parts lighter, more olivaceous brown, and the throat,

* Two specimens measure; wing 80-82; tail 79; bill 19-195 mm.

BIRDS OF WESTERN COLOMBIA. 1089

foreneck, &c. of a much paler, orange-rufous. This variation is
well shown by our series of the nearly allied L. p. lawrencii
Lawr.* from La Vijagua, Eastern Costa Rica fT.

L. p. pheeocephalus is confined to the lowlands of W. Ecuador
and W. Colombia.

6. THRYOPHILUS NIGRICAPILLUS scHoTri Baird.

[Lhryothorus nigricapillus Sclater, P. Z. 8. 1860, p. 84 (1860.—
Nanegal, W. Ecuador). |

Thryophilus schottii Baird, Review Am. B. 1, p. 133 (1864.—
Truando R., Colombia).

Thryothorus nigricapillus (nec Sclater) Cassin, Proc. Acad. N.
Sei. Philad. 1860, p. 195 (River Truando).

Thryophilus nigricapillus (nec Sclater) Sclater & Salvin, P. Z.5
1879, p. 493 (Remedios, Antioquia).

No. 2585. gd ad. Condoto: 2.iv.09.—Wing 67; tail 51; bill
18 min.

Nos! 2181) 258i 2622) OO "ad! Sip] 2s 08)-* Condoto:
20.111., 22.iv.09.— Wing 63-65; tail 46-48 ; bill 17 mm.

Nos. 2618, 2621. 9 2 juv. Condoto: 21,22.iv.09.— Wing 633,
65; tail 46, 48; bill 15 mm.

“Tris brown, feet dark grey, bill black, mandible grey in adults,
yellow in young birds.”

These specimens differ from a good series of 7. n. nigricapillus
from Western Heuador in having the black cross-bands of the
lower parts much broader, more regular as well as more closely
set, especially along the middle Ag the belly. Moreover, chin
and. throat ere distinctly, though narrowly, banded with. blackish,
while they are uniform white in the Ecuadorian form. Even
young birds can easily be distinguished by these characters. In
the coloration of the lower parts T. n. schottit approaches ~
T. sennibadius Salv., of Chiriqui and Costa Rica, but has the top
of the head black like 7’. n. nigricapillus. Its range is evidently
restricted to Western Colombia, from the R. Tr uando south to
the San Juan district. -Like its southern representative, it
exclusively lives in the hot, low country.

. THRYOPHILUS LEUCOPOGON Salvad. & Festa.

Phnyophilus leucopogon Salvadori & Festa, Boll. Mus. Zool.
Torino, xiv. no. 357, p. 6 (1899.—Rio Peripa, W. er
Hellmayr, Journ. f. Orn. 1903, p. 534 (San Javier, N.W.
Ecuador).

No. 2440. g ad. Névita: 28.xii.08.—Wing 58; tail 38; bill

163 mm.

* Cyphorinus lawrencii Lawrence (ex Sclater, MS.), Ann. Lye. N. H. N.Y. viii. p.5
(May 1863,—based on C. cantans (nec Gmelin) Lawrence, 1. ¢. vil. 1861, p. 293.—
Panama Railroad).

+ The presence of dusky bars on the primary coverts upon which the late
Dr. Sharpe Jaid much stress is a purely individual character. Five birds from
W. Ecuador al/ have distinct blackish bars, as also the female trom Juntas.

1090 MR. CG. E. HELLMAYR ON THE

“Tris dark red, feet light grey, maxilla black, mandible grey.”

This specimen agrees perfectly with others from Western
Keuador. In addition to the one from San Javier in the Vienna
Museum mentioned by me, /. c., I have since examined four more
Specimens from N.W. Ecuador in the Tring Museum; one male
and two females also from S. Javier, and a male from Lita,
3000 ft. The wing measures 54-58, the tail 33-37 mm.

T’. leucopogon, though a very well-marked species, is nearest to
T. thoracicus Salv., of Costa Rica, which it closely resembles on
the upper parts, but differs mim the uniform dull ochreous brown
colour of the lower surface, with the exception of the chin and
upper throat which are white with slight blackish edgings, while
in 7’. thoracicus all the throat and breast feathers are white,
broadly margined with black laterally.

T. leucopogon is known only from the coast-belt of Western
Ecuador and Colombia.

8. HenicorRHINA INORNATA Hellm.

Henicorhina imornata Hellmayr, Journ. f. Ornith. l. p. 528
(1903.—Lita, N.W. Ecuador).

No. 2135. 2 ad. Sipi, 30.ix.08.—Wing 55; tail 2743; bill
16 mm.

‘‘ Tris dark brown, feet and bill black.”

This bird is practically identical with the series, including the
type, from N.W. Ecuador, in the Munich Museum. The upper
parts are of the same bright chestnut-rufous hue, the sides of the
breast deep smoky grey, the flanks dark rufous brown, and the
base of the lower mandible is clear yellowish white. H. inornata
is known only from Western Colombia and N.W. Kcuador, where
it inhabits the forests of the humid lowlands as well as the lower
slopes, up to 3000 feet (Lita).

[In the mountains bordering the Cauca Valley it is repre-
sented by another species which Mr. Bangs * has lately separated
as H. leucosticta eucharis, but which I cannot satisfactorily
distinguish from H. 1. prostheleuca, of Chiriqui and Eastern Costa
Rica. Having before me two fine specimens t, I can positively
state that it has nothing to do with H. lewcosticta of Guiana and
Kast Venezuela. Mr. Bangs was apparently misled by a black-
crowned male, but this character is of very little importance in
view of the fact that specimens of prosthelewca in worn plumage
often have the crown nearly uniform black, the brown tips to the
feathers having disappeared through abrasion. Moreover, the two
skins from Primavera have the feathers of the pileum broadly
tipped with umber-brown and the back ete. dull russet- brown,
exactly as in prostheleuca. While a larger series from Colombia
might ultimately reveal some slight differences, the evidence at
hand is not in favour of the southern form being separable. |

* Proce. Biol. Soc. Wash. xxiii. p. 74 (1910.—Pavas, W. Colombia, 4400 feet).
+ & & trom Primavera, Cauca R., 5100 feet, Raap coll., Tring Museum.

BIRDS OF WESTERN COLOMBIA. 1091

9. SEIURUS NOVEBORACENSIS NOVEBORACENSIS Gin.

Motacilla noveboracensis Gmelin, Syst. Nat. 1, i. p. 958 (1789
—ex Waubenton, Pl. Enl. 752. fig. 2: Louisiana).

Siurus noveboracensis Sclater & Salvin, P. Z. 8. 1879, p. 493
(Concordia, Medellin).

Nos. 2131, 2153, 2171. 3b @ ad., (sex not determined). Sipi:
TOR Box Ole,

“Tris dark brown, feet brown, maxilla black, mandible brown.”

These birds apparently belong to typical noveboracensis. Quite
similar specimens we have from Ecuador and Bogota collections.

A common winter visitor to Colombia.

10. Mwnioriuta varia Linn.

Motacilla varia Linneus, Syst. Nat. xii. 1, p. 333 (1766—ex
Brisson & Sloane: Jamaica and San Domingo).

Miiotilta varia Sclater & Salvin, P. Z. 8. 1879, p. 493 (Con-
cordia etc.); Berlepsch, Journ. f. Orn. 1884, p. 282 (Bucara-
manga).

Nos. 2807, 2855. § 2. Pueblo Rico: 27.x.; Loma Hermosa:
Daexa)o: :

“ Tris and feet dark brown, maxilla black, mandible white.”

A common winter visitor to Colombia.

11. DeNpDROICA CASTANEA Wilson.

Sylvia castanea Wilson, Amer. Orn. il. p. 97, pl. 14. f. 4 (1810.
—Pennsylvania).

Dendreca castanea Sclater & Salvin, P. Z.S. 1879, p. 494
(Remedios); Berlepsch, Journ. f. Orn, 1884, p. 282 (Bucara-
manga). —

Nos. 2317, 2331, 2381. § dg juv. Novita: 20,24.xi., 9.xii.08.

“Tris dark brown, feet grey, maxilla black, mandible grey.”

12. DENDROICA ZSYIVA ZSTIVA Gin.

Motacilla cestiva Gmelin, Syst. Nat. 1, ii. p. 996 (1789—ex
Brisson & Daubenton, Pl. Enl. 58, fig. 2: Canada),

Dendreca estiva, Sclater & Salvin, l. c. p. 494 (Medellin) ;
Berlepsch, |. c. p. 282 (Bucaramanga).

Nos. 2593, 2136. g ad., ¢ juv. Condoto: 12.ix.; Sipi:
30.ix.08.

“Tris dark brown, feet light brown, maxilla black, mandible

grey.”

13. BASILEUTERUS TRISTRIATUS TRISTRIATUS Tsch.

Myiodioctes tristriatus Tschudi in Arch. f. Naturg. 10, 1. p. 283
(1844.—Peru); cfr. Berlepsch & Hellmayr, Journ. f. Orn. 1905,
p. @ (erit.).

Basileuterus tristriatus Berlepsch, Journ, f. Orn, 1884, p. 283
(Bucaramang’).

1092 MR. C. E. HELLMAYR ON THE

SL. auricularis Sharpe, Cat. B. Brit. Mus. x. p. 386 (1885.—
Bogota; Pallatanga, W. Ecuador; Simacu, Bolivia; no type
specified).

B. melanotis dedalus Bangs, Proc. Biol. Soc. Wash, xxi.
p- 160 (1908.—San Antonio, Rio Cali, W. Colombia, 5800 ft.).

No. 3748. g ad. Pueblo Rico, 5200 ft., 10.1x.09.— Wing 65 ;
tail 57; bill 103 mm.

Nos. 3749, 3751. 2 (@)ad., Q ad. Pueblo Rico: 10,11.1x.09.—
Wing 64, 613; tail 60, 56; bill 10 mm.

“ Tris dark brown, feet greyish yellow, maxilla black, mandible
grey.”

These specimens agree with a series from Bogota, Ecuador, and
Peru, some of which I had previously compared and found
identical with Tschudi’s type kindly forwarded by the authorities
of the Neuchatel Museum. Iam unable to discover any constant
character on which to separate the Colombian and Ecuadorian
birds from typical Peruvian skins, 6. auwricularis Sharpe being
based on purely individual variations, such as the colour of
the pale crown stripe etc. When compared with B. tristriatus
melanotis Lawr. *, of Costa Rica and Chiriqui, the three examples
from Pueblo Rico differ exactly as indicated by Mr. Bangs,
viz., much brighter green back, deeper (about maize) yellow
underparts, and large, blackish loral spot. However, in all these
points they are absolutely similar to typical tristriatus, to which
Mr. Bangs, evidently misled by Sharpe’s key, does not make
any reference at all. B. t. tristriatus had already been recorded
from Bucaramanga, N. Colombia, by Count Berlepsch.

While I cannot admit the distinctness of the Colombian birds—
for which, moreover, Sharpe’s term auricularis would be an earlier
name—the inhabitants of the mountains of Venezuela constitute
a fairly well-defined race, B. ¢. meridanus Sharpe 7, recognisable
by the reduction or absence of the black loral and auricular spots.

We have in the Munich Museum, three adults from Merida,
and five males and one female from the Cumbra de Valencia ;
and, at Tring, I have examined two skins from Caripé, State of
Cumana, and one adult from Buearito, Tocuyo (Mocquerys).

14. BASILEUTERUS FULVICAUDA SEMICERVINUS Scl.

[ Muscicapa fulvicauda Spix, Av. Bras. 11. p. 20, pl. xxviii. fig. 2
(1825—no locality) |.

Basileuterus senurcervinus Sclater, P. Z. 5. 1860, p. 84 (1860.—
Nanegal, W. Ecuador); Sclater & Salvin, P. Z. 8. 1879, p. 494
(Remedios, Neche); Berlepsch, Journ. f. Orn. 1884, p. 284
(Bucaramanga).

No. 613. Q ad. Juntas, Rio Dagua, 1000ft.: 1.viii.07.—
Wing 60; tail 49; bill 123 mm.

* B. melanotis Lawrence, Ann. Lyc. N. H. N.Y. ix. p. 95 (1868.—Cervantes, Costa

Rica). :
+ B. meridanus Sharpe, Cat. B. Brit. Mus x. p. 3887 (1885.—Merida, Venezuela).

BIRDS OF WESTERN COLOMBIA. ~ 1093

No. 2170. Q ad. Rio Garrapatas, Sipi, 150 ft.: .8.x.08.—
Wing 60; tail 48; bill 12 mm.

“Tris dark brown, feet light brown, bill black.”

These, as well as several other specimens from W. Colombia
in the Tring Museum (Raap and Palmer coll.), are practically
identical with topotypical skins from Nanegal, W. Ecuador. A
series from N.W. Ecuador is not different either.

15, SETOPHAGA RUTICILLA Linn.

Motacilla ruticilla Linneus, Syst. Nat. x. p. 186 (1758—ex
Catesby : Carolina).

Setophaga ruticilla Sclater & Salvin, 1. c. p. 494 (Concordia,
Medellin) ; Berlepsch, 1. c. p. 284 (Bucaramanga).

No. 2392. gad. Névita: 11.x11.09.

No. 3775, —. g gad. Pueblo Rico: xi.09; Siaté: 25.xi.09.

“Tris dark brown, feet black, maxilla black, mandible
brown.”

A common winter visitor to the north-western States of Bone
America.

16. PRoGNE CHALYBEA CHALYBEA Gm.

Hirundo chalybea Gmelin, Syst. Nat. 1, ii. p. 1026 (1789—
ex Brisson and Buffon: Ca emo

Progne chalybea Sclater & Salvin, |. c. p. 495 (Remedios).

Nos. 2356, 2360. ¢ g ad. Ndévita: 30.x1., 1.xii.08.—Wing 128,
125% tail 69; bill 12, 11 mm.

“Tris, feet, and bill black.”

Agreeing with examples from Venezuela, Bogota, and Cayenne,

17. SrELGrDOPrERYX RUFICOLLIS UROPYGIALIS Lawr.

[Hirundo ruficollis Vieillot, Nouv. Dict. xiv.. p. 523 (1817.—
“ Brésil ”’).|

Cotyle uropygialis Lawrence, Ibis, v. p. 181 (1863.— Panama).

Stelgidopteryx uropygialis Sclater & Salvin, |. c. p. 496
(Remedios); Berlepsch, J. f. Orn. 1884, p. 285 (Bucaramanga).

Nos. 2106, 2107. ¢ Q ad. Sipi: 22.ix.08.—Wing 107, 92;
tail 56, 47 mm.

‘“‘ Tris dark brown, feet and bill black.”

Agreeing with specimens from Chiriqui and Costa. Rica. Cfr.
Nov. Zool. xiii. 1906, p. 13.

18. DIGLOSSOPIS CHRULESCENS CARULESCENS Scl.

Diglossopis cerulescens Sclater, Ann. Mag. N. Hist. (2) xvii.
p- 467 (1856.—Caraceas, in Venezuela) ; Sclater & Salvin, P. Z.S.
1879, p. 496 (Santa Elena).

Dig cerulescens Berlepsch, Journ. f. Orn. 1884, p. 286

(Bucaramanga).

IEROC ACO, SOUR ING: IRON, re!

(1094 MR. C. BE. HELLMAYR ON THE

No. —. gad. Tatama, 2794 ft.: 12.x.09.—Wing (2; tail 58;
bill 13 mm.
‘Tris reddish brown, feet dark grey, bill black.”

Agrees with Bogota skins, whereas specimens from Merida are

a darker and more uniform bluish colour below with scarcely
any greyish admixture on the vent. Topotypical birds from
Caraccas are not available for comparison.

D. c. cerulescens is known only from the mountains of Colombia
and Venezuela. In Peru it is replaced by the nearly allied
D. c. pallida Berl. & Stolzm.*

19. DACNIS VENUSTA FULIGINATA Bangs.

'Dacnis venusta Lawrence, Ann. Lyc. N. H. N.Y. vii. p. 464
(1862.—Panama Railroad) |

D. venusta fuliginata Bangs, Proc. Biol. Soc. Wash. xxi. p. 160
(1908.—Jiménez, W. Colombia).

D. venusta (nec Lawrence), Sclater & Salvin, P. Z. 8. 1879,
p- 497 (Remedios).

No. 2215. gad. Noanama: 22.x.08.—Wing 62; tail 39;
bill 11 mm.

Nos. 2243, 2244. § gad. Novita: 7.x.08.—Wing 643, 65;
tail 403, 41; bill 10, 11 mm.

No. “385. gad. Jiménez, 1600ft.: 12.vi.07.—Wing 63;
tail 40; bill 103 mm.

No. —. d a Rio Dagua: 8.vi.95. Rosenberg coll.— Wing 63 ;
tail 403; bill 10 mm.

No. —. Q ad. Rio Dagua: 10.vi1.95. Rosenberg coll. Wing
64; tail 42; bill 11 mm.

‘“‘ Tris crimson, bill and feet black.”

In addition, I have examined two males from Jiménez (June
14, 28), in Mr. Rosenberg’s possession, D. v. fuliginata is a
perfectly good form. Compared with the series from Panama,
Chiriqui and Costa Rica in the Munich Museum, the males sent
by Mr. Palmer differ in having the underparts much darker,
deep black with hardly a trace of the greenish hue so conspicuous
in the Central American birds. The female is smaller tT, and
rather deeper ochraceous buff on the lower belly and under tail-
coverts than those from more northern localities. The bill, in
the southern race, is constantly shorter. Birds from Costa Rica
and Chiriqui have much longer wings and tail, while those from
Panama, in size, approach D. v. fuliginata.

Hight adult males from Chiriqui and Costa Rica measure :
wing 67-70; tail 43-45; bill 12-13 mm.

Three adult males from Panama (Railroad) measure : wing 633—
66; tail 42-43; bill 12-13 mm.

* P, Z.S. 1896, p. 334 (Chachapoyos, N. Peru ; Garita del Sol, C. Peru).
+ Three adult females SNH! Chiriqui measure: wing 65-67; tail 43-443; bill
—13 mm.

BIRDS OF WESTERN COLOMBIA. 1095

20. DAGNIS CAYANA CG@REBICOLOR Scl.

[ Motacilla cayana Linneus, Syst. Nat. xu. 1, p. 336 (1766—
ex Brisson: Cayenne; excl. Hernandez—Mexico)}.

Dacnis cerebicolor Sclater, Contrib. to Ornith. 1851, pp. 106—
12 (1851 (?)—“ New Granada?” sc. Bogotd, cfr. Sclater, Cat.
Coll, Amer. B., 1862, p. 51); Sclater & Salvin, P.Z.S. 1879,
p- 497 (Remedios— 3 imm., ‘‘not quite so bright as specimens
from Bogota”); Hartert, Nov. Zool. v. p. 481 (Paramba, N.W.
Keuador).

D. cayana (errore) Sclater & Salvin, 1. ec. p. 496 (Remedios).

Nos. 2006, 2205, 2214. ¢ gad. Noanama: 24.viii., 20.22.x.08.
—Wing 61-644; tail 424-434; bill 11-114 mm.

Nos. 2319, 2348, 2187. ¢ gad. Névita: 21,28.xi.; Sipi: 13.x.
08.—Wing 61-63; tail 424-44; bill 11-112 mm.

No. 2657. gad. Tadd: 21.v.09.—Wing 63; tail 42; bill
12 mm.

No. 2007. @ ad. Noanama: 24.viii.08.—Wing 59; tail 42;
bill 12 mm.

Nos. 2320, 2374, 2382. 9 9 ad. Né6vita: 21.xi., 7, 9.xii.08.—
Wing 59-62 ; tail 40-43; bill 104-12 mm.

Nos. 2188, 2073. 9 Q ad. Sipi: 10i1x., 13.x.08.—Wing 63,
61; tail 43, 424; bill 11 mm.

Nos. 2631, 2669. 9 2 imm. Tadé: 3, 27.v.09.—Wing 60, 62;
tail 43, 44; bill 11, 114 mm.

Nos. 507, 508, 518, 541, 564, 570, 647,688. ¢ gad. Jiménez,
1600 ft., 28.vi., 12, 13, 18, 23, 24.vii., 9.vi1i.07.—Wing 634-67;
tail 43-47 ; bill 12-13 mm.

No. —. g ad. Rio Oscuro, W. Cordillera, 3000 ft., June 1898,
Batty coll.— Wing 68; tail 49; bill 123 mm.

-Nos, 384, 400, 401, 486, 577, 642,652. 9 9. Jiménez, June to
Aug.— Wing 60-63; tail 42-44; bill 113-12 mm,

“Tris dark red in males, dark brown in females, feet pink,
bill black.”

This series is rather puzzling. The males show a wide amount
of individual variation, every shade of blue between the ultra-
marine of D. c. napaea and the deep purplish blue of D. c. cwre-
bicolor (of Bogota) being represented. The lightest specimen,
no. 2006, from Noanama has the blue portions of the plumage of
exactly the same tone, and the mantle as well as the gular patch
as dull (greenish black) as topotypical Santa Marta skins, from
which it only differs by its smaller size and shorter bill. Next
come no. 2214 (Noanama), no. 564 (Jiménez), and the male froin
Rio Oscuro, which are just a shade darker, more cobalt, with the
black of the mantle and throat deeper, less obscured by dull
greenish tips. Then follow ten specimens which have the plumage
more or less tinged with purple. Finally, three birds, nos. 2187
(Sipi), 508 and 541 (Jiménez), and a fourth from Paramba, N.W.
Keuador, are quite as bright and deep purplish blue as Bogota
skins.

74*

1096 MR. C. E. HELLMAYR ON THE

In coloration, there is no constant difference between the birds
from the foot-hills (Noanama, Névita, Tad6, Sipi) and those
from the more elevated districts (Jiménez, 1600 to 2900 ft. ; Rio
Oscuro, 3000 ft.). The former are, however, on the average, smaller,
and have a shorter, slenderer bill, while the males from Jiménez,
etc., in dimensions, agree with skins from Bogota and Buca-
ramanga.

The females also are rather variable. Most of them have the
top and sides of the head turquoise-blue, scarcely darker than in
the female of D.c. wltramarina from Panama, while in two or
three these parts are of the same dark ultramarine-blue colour as
in Bogota skins of D. c. cwrebicolor.

From what I have said above, it is evident that the majority
of the West Colombian birds are not typical ewrebicolor ; but in
view of their great individual variation, which completely connects
napaea, of Santa Marta, and cwrebicolor, of the Kastern Cordillera,
it seems inadvisable to separate them subspecifically since the new
“form” would mainly consist of ‘‘ intermediates.” One thing,
however, results from the study of that series, viz., that both
napaea and cerebicolor are merely geographical races of D. cayana,
the passage being formed by D.c. ultramarina. 'Thus, we have
the following forms :—

(a) D.c. cayana Linn. Eastern South America from Southern
Brazil to Trinidad, Guiana and Venezuela.

(b) D. ¢. glaucogularis Berl. & Stolzm.* From the eastern
slopes of the Colombian Andes south through Peru to
Eastern Bolivia and Western Brazil (Mattogrosso).

(c) D. c. callaina Bangs+. Chiriqui and 8.W. Costa Rica
(Pozo Azul ete.).

(d) D.c. ultramarina Lawr.t Isthmus of Panama, Eastern
Costa Rica, H. Nicaragua.

(e) D. c. napaea Bangs§. Santa Marta District, N. Colombia.

(f) D. c. cerebicolor Scl. Andes of Colombia (Western and
Eastern Cordillera) and N.W. Ecuador. Typical in the
Eastern Cordillera.

On some future occasion I hope to give more details about the
variation and geographical distribution of the various races.

21. CHLOROPHANES SPIZA EXSUL Berl. & Tacz. (2)

[Motacilla spiza Linneus, Syst. Nat. x. p. 188 (1758—ex
Edwards: Surinam). |

Chlorophanes spiza exsul Berlepsch & Taczanowski, P. Z.5S.
1883, p. 543 (1884—Chimbo, 8.W. Keuador).

* P. Z.S. 1896, p. 336 (Central Peru: La Gloria, La Merced).

+ Proc. Biol. Soc. Wash. xviii. p. 154 (1905.—Divala, Chiriqui).
Proc. Acad. N. Sci. Philad. 1864, p. 106 (Isthmus of Panama).
Proc. Biol. Soc. Wash. xii. p. 143 (1898.—Santa Marta),

BIRDS OF WESTERN COLOMBIA. 1097

C. atricapilla (errore) Sclater & Salvin, P. Z. 8. 1879, p. 497
(Concordia, Remedios). -

Nos. 2211, 2248, 2385. ¢ gad. Noanama, 22.x.;- Novita:
8.xi., 10.x11.08,— Wing 69 ; tail 45-48; bill 13-133 mm.

Nos. 2212, 2333. ¢g juv., Qad. Novita: 24.xi.; Noanama :
22.x.08.— Wing 68, 65; tail 45; bill 133 mm.

“Tris crimson, feet green, maxilla black, mandible yellow.”

The adult males agree with typical specimens from West
Ecuador in size and in smallness of the bill; but the coloration
is more bluish, especially below, though much less so than in the
Upper Amazonian race, C. s. cerulescens Cass.* Birds from
Chiriqui and Costa Rica (El General de Terraba, Carrillo, La
Vijagua, ete.) are again more greenish, have longer wings (70—
74 mm.), and a longer, more robust bill (143-163 mm.). They
are unquestionably subspecifically distinct. A larger series might
perhaps enable us to separate the West Colombian birds, but for
the present I leave them with C.s. exsul. According to this view,
C. s. exsul ranges from 8.W. Ecuador north to the Pacific slopes
of Colombia.

22. CYANERPES CHRULEA MICRORHYNCHA Berl.

[Certhia cerulea Linneus, Syst. Nat. x. p. 118 (1758—ex
Edwards: Surinam). |

Cereba cerulea microrhyncha Berlepsch, Journ. f. Orn. xxxil.
p. 287 (1884—part. : type ex Bucaramanga, N. Colombia).

Cereba cerulea (nec Linneus) Sclater & Salvin, P. Z. S.
1879, p. 497 (Remedios, Medellin).

Nos. 2257, 2262. ¢ gad. Novita: 10,11:xi1.08.—Wing 55,
534 ; tail 29, 273; bill 18 mm. 3

No. 1961. g ad. San Joaquim, Bahia del Choeé, 3.vi1.08.—
Wing 54; tail 28; bill 173 mm.

No. 2241. g ad. Rio Cajén: 5.xi.08.—Wing 55; tail 28;
bill 173 mm.

No. 2238. ¢ juv. Cajon: 2:x1.08.

Nos. 2242, 2349. 92 Qad. Rio Cajén: 5.xi.08; Noévita:
28.xi.08.— Wing 55; tail 27; bill 17 mm.

“ Tris dark brown (¢ ¢), black (2), bill black, feet wax-yellow
(3), Seem (2)

The adult males agree in the shortness of the bill, and in the
pale azure-blue colour of the cheeks and anterior portion of the
crown, with a number of Bogota skins, but have slightly shorter
wings and tail. Two adult males from Pozuzo (prov. Huanuco,
Peru) are equally small while the bill is even a little shorter.
The difference in size is, however, not likely to be constant, since
a single male from San Augustin, N. Bolivia (3500 feet), has
wings, tail, and bill fully as long as Colombian examples. There

* Chlorophanes cerulescens Cassin, Proc. Acad. N. Sc. Philad. Nov. 1864, p. 268
(1865.—Yuracares, N.E. Bolivia).

1098 MR. GC. E. HELLMAYR ON THE:

can be no question that C.c. microrhyncha is a perfectly valid
form. Cfr. my remarks in Nov, Zool. xii. 1905, p. 9, and J. ¢. xiv.
1907, pp. 42-43.

Its range extends from the mountains of Colombia south to
Northern Bolivia.

23. C@REBA CHLOROPYGA MEXICANA Scl.

[Certhiola chloropyga Cabanis; Mus. Hein. i. p. 97 (1850.—
Bahia). ]

Certhiola mexicana Sclater, P. Z. S. 1856, p. 286 (Jan. 1857.—
S. Mexico, probably Vera Cruz); Sclater & Salvin, P. Z. 8.
1879, p. 497 (Remedios, Medellin).

No. 2822. 9 ad. Pueblo Rico, 5200 ft. : 2.xi.09.— Wing 523 ;
tail 32; bill 12 mm.

“Tris dark brown, feet dark grey, bill black.”

Identical with skins from Bogota and Western Ecuador which
I am unable to separate from others taken in Mexico and
Costa Rica. ;

Birds from Bucaramanga form the transition to C. c. luteola
Cab. Cfr. Berlepsch, Journ. f. Orn. 1884, p. 288.

24. ORYZOBORUS FUNEREUS £THIOPS Scl.

[Oryzoborus funereus Sclater, P. Z.S. 1859, p. 378 (1860.—
Oaxaca, Mexico). |

Oryzoborus cwthiops Sclater, P. Z. 5. 1860, p. 88 (1860.—
Nanegal, W. Ecuador).

No. 1988. g ad. Mouth of Calima, San Juan River, 13.vii.08.
—Wing 55; tail 51; bill 12 mm.

‘‘Tris dark brown, feet and bill black.”

This bird agrees, in size and colour, with a series from Western
Ecuador in the Munich Museum. Specimens from Guatemala,
which I take to be typical O. f. funereus, have longer wings
(60 mm.) and tail (54 mm.), a slightly larger bill, and the females
are much brighter cinnamon underneath. These slight differences
might disappear on comparison of a larger series of Central
American skins, but, for the present, I am unwilling to unite the
two forms without further evidence. O./. ethiops, as defined
here, ranges over the forest district of Western Colombia and
Ecuador. Specimens from Antioquia * and Santa Marta which
I have not seen may also be referable to the southern race.

25. SPOROPHILA OPHTHALMICA Scl.

Spermophila ophthalmica Sclater, P. Z. 8. 1860, p. 276 (end
of 1860—Babahoyo, W. Ecuador); Sharpe, Cat. Birds, xii.
p- 120 (Chocé Bay).

Nos. 1979, 1991. ¢ gad. Mouth of Calima, 13.vili.08;

* O. funereus Scl, & Salvin, P. Z. S. 1879, p. 506 (Medellin).

BIRDS OF WESTERN COLOMBIA. 1099

Guineo, R. Calima: 7.viii.08.—Wing 54-55; tail 45, 47;
bill 10 mm.

No. 2134. Q ad. Sipi: 30.ix.08.—Wing 53; tail 43;
bill 10 mm.

‘‘ Tris dark brown, feet and bill black.”

One of the males (no. 1979: Guineo) agrees with West
Ecuadorian skins, the throat being white with a small black spot
at the base of the mandibular rami. In the other specimen
(no. 1991) the feathers of the upper throat are black, tipped with
white. Both have the black breast-band rather wider than the
large majority of Ecuador skins, though one or two of the latter
hardly differ in that respect.

The male from the mouth of the Calima (no. 1991) approaches
very closely black-throated males of S. aurita, and can only be
distinguished by the white tips to the feathers of the throat !

S. ophthalmica is no doubt merely a southern race of S. aurita™,
but until the status of this puzzling “ species” is better understood,
it would be unwise to employ a trinomial appellation. Mr.
Ridgway’s remarks (in Bull. U.S. Mus. no. 50, Part 1, 1901,
p. 573) should be carefully consulted.

S. ophthalmica is restricted to W. Ecuador and 8.W. Colombia.

96. TERSINA T VIRIDIS OCCIDENTALIS Scl.

[Hirundo viridis Mliger, Prodr. Syst. Mamm. & Av. p. 229
(1811—based on “ L’Hirondelle verte,’ Temminck, Cat. Syst.,
1807, p. 245, no. 986: Brasilia;= ©); cfr. Allen, Bull. Amer.
Mus. N. Y. ii. 1889, p. 70.]

Procnias occidentalis Sclater, P. Z.S. 1854, p. 249 (Apr. 1855.
—-“‘ Nova Grenada ”).

P. tersa (nec Linneus) Sclater & Salvin, P. Z. 8, 1879, p. AQT
(Remedios, Antioquia).

P. cerulea occidentalis Berlepsch, Journ. f. Orn. 1884, p. 288
(Bucaramanga).

No. 2035. gad. Noanama: 29.viii.08.—Wing 87; tail 56 ;
bill 9 mm.

Nos. 2404-6, 2412. gdad. Noévita: 16, 18.xii.08—Wing
81-85 ; tail 52-55: bill 9-10 mm.

‘‘ Tris brown, feet and bill black.”

The series agrees, in size and coloration, with other specimens
from Bogota, Eastern Ecuador, Venezuela (Cumana), etc. Cr.
my remarks in Nov. Zool. xvii. 1910, p.271. Birds from Eastern
Brazil (Bahia, Rio) are much larger, and the males have the
plumage of a decidedly paler, more greenish blue (nile-blue).

* Not having seen white-throated males (=hicksii Lawr.) Iam unable to say
in what way they differ from S. ophthabmica, but, judging from published de-
scriptions, the only character of the northern form appears to consist in the rather
wider black band across the chest.

+ For change of generic name see Ridgway, Bull. U. S. Mus. no. 40, pt. iv. p. €89,
footnote b

1100 ; MR. GC. E. HELLMAYR ON THE

27. EuPHoNIA FruLyrcriIssa Sel. [a subsp. ®].

Euphonia fulvicrissa Sclater, P. Z.S. 1856, p. 276 (Jan.
1857.— “Santa Martha in New Grenada”); Cassin, Proc.
Acad. N. Sci. Philad. 1860, p. 143 (Falls of the Truando) ;
Selater & Salvin, P. Z. S. 1879, p. 498 (Remedios, Neche:
Antioquia); Berlepsch, Journ. f. Orn, 1884, p. 288 (Bucara-
manga).

No. 2036. g ad. Noanama, 29.viii.08.—Wing 51; tail 29;
bill 82 mm.

No. 2530. Q ad. El Tigre, 10.11.09.—Wing 50; tail 28;
bill 8 mm.

“Tris dark brown, feet black, bill black, lower mandible blue.”

The adult male from Noanama and another, picked out from a

set of Bogota skins, in the Munich Museum are exactly inter-
mediate between specimens of H. fulvicrissa, from Panama and
Chiriqui, and #. f. purpurascens Hart.*, from N.W. Ecuador.
The back, upper wing-coverts, sides of the head and throat are
glossy metallic steel-blue in the Colombian skins, dull greenish
‘blue or bottle-green in those from Central America, and bright
purple-blue, inclining to violet, in purpurascens. Two skins of
the last named form? have very little, or no white at all, on the
outermost rectrix, while in the specimens from Chiriqui,
Panama, and Bogota a.large portion of the inner web is white.
As, however, the male from Noanama agrees, in that respect, with
purpurascens, much importance cannot be laid upon this
character.
_ The type of Z. fulvicrissa in the British Museum, said to be
from Santa Marta, where, however, recent travellers did not meet
with the species, is to be examined in order to ascertain whether
it belongs to the Colombian ov Veraguan race.

28. KuPHONIA XANTHOGASTER CHOCOENSIS Hellm.

[Euphone sxanthogaster Sundevall, Vetenskaps Akad. Hand.
1833, p. 310, pl. 10. fig. 1 (= ¢ ad.) (1834.—* Brazil”).

Euphonia wanthogaster chocoensis Hellmayr, Rey. Frang. @Orn.
no. 22, p. 23 (1911.—Rio Cajén, W. Colombia).

No. 2015. gad. Noanama: 26.vili.0@2—Wing 61; tail 354;
bill 74 mm.

Nos. 2235, 22367. g¢ g ad. Rio Cajén: 3.xi.08— Wing 58,
D9; tail 33, 34; bill 8 mm.

Nos. 2086, 2176. 9 9 ad. Sipi: 16.ix., .10.x.08.— Wing 56,
57; tail 30, 31; bill 73-8 mm.

“Tris black (¢), dark brown ( @ ), feet dark grey (3), grey (2),
bill black, mandible blue.”

_ * Novit. Zool. viii. p. 377 (1901.—Pambilar, N.W. Ecuador; type in Tring
Museum).

+ The type from Pambilar, and anotuer from San Javier, both in the Tring
Museum.

t Type of subspecies.

BIRDS OF WESTERN COLOMBIA, 1101

' This new subspecies is most nearly allied to typical E. wantho-
gaster, from: Brazil, Peru and Ecuador, but differs in its generally
smaller size, especially shorter and, smaller bill. The males, too,
have the underparts much clearer chrome-yellow, without any
trace of the orange tinge which is always well pronounced along
the middle line in the eastern race; the females also may be
distinguished by the nearly pure cinereous instead of buffy grey
colour of the throat and breast, and by having the middle of the
abdomen much paler buffy. Forehead and crown, in the males,
are clear chrome-yellow, as in average Spee ines of wanthogaster.

E. xanthogaster brevirostris Bonap.*, from Bogota, is much
larger (wing 64-65; tail 38-40; bill 9 ‘mm. ), has the foreneck
and middle of the belly tinged with orange, and the occiput of a
deep brownish orange colour. 4. «. brunneifrons Chapm. t, from
South-eastern Peru, is of the same size as brevirostris, but the
crown is still darker, dull orange rufous.

The new form has been compared with large series of the
various races. Besides the thirteen specimens in the Munich
Museum, I had before me twelve from the Berlepsch Collection,
and three belonging to the Vienna Museum.

29. CHLOROCHRYSA NITIDISSIMA Scl.

Chlorochrysa nitidissima Sclater, P. Z. 5. 1873, p. 728 (1874. —
Antioquia); idem, Ibis, 1875, p. 466, pl. x.; Sclater & Salvin,
P. ZS. 1879, p. 498 (Jerico, Antioquia).

No. 3757. 3S imm. Siat6, Rio Siaté: 16.1x.09.—Wing 67 ;
tail 44°; bill 12 mm.

No, 3745. 9 imm. Pueblo Rico: 9.1x.09.—Wing 67; tail 43;
bill 13 mm.

“Tris brown, feet grey, bill black.”

This species was discovered by the late T. K. Salmon at Jerico,
south of Medellin, on the left side of the Cauca River, in the
Western Cordillera of Colombia. Dr. Sclater described and
figured the adult male. Specimens transmitted by the late J.
H. Batty from Riolima (Cauca Valley) agree perfectly with the
or iginal description.

The immature male and female differ in lacking the lemon-
yellow interscapulium (which is mainly green with but a faint
yellowish tinge) and in having the middle of the belly dull grey
instead of black. Moreover, the general colour is not so bluish.

C. nitidissima is apparently restricted to the Western
Cordillera of Colombia, and found only at altitudes above
5000 feet.

30. CALOSPIZA JOHANNA Dalmas.

Calliste johanne Dalmas, Bull. B. O.C. xi. p. 36 (1900.—E]

* Huphonia brevirostris Bonaparte, Rev. Mag. Zool. (2) ili. p, 136 (1851; ex
Columbia).

HE Euphonia wanthogastra Bysnanmnesseaawas Chapman, Bull. Amer. Mus, N. H. xiy.
226 (1901.—Inca Mines, 8.E. Peru).

1102 MR. C. E. HELLMAYR ON THE

Paillon, near Buenaventura, Chocé, W. Colombia) ; Sclater, Ibis,
1901, p. 597, pl. xii. fig. 2 (Paramba, N.W. Ecuador).

No. 2698. gad. Tadé, 7.vi.09.—Wing 73; tail 50; bill
105 mm.

No. 2388. 9 ad. Novita: 10.xii.08.—Wing 67; tail 46;
bill 10 mm.

No. 2589. ¢ juv. Condoto: 10.iv.09.—Wing 714; tail 49;
bill 9 mm.

“Tris dark brown, feet dark grey, bill black.”

Two more specimens, a female and an immature bird, sex not
determined, from Tadé, June 1909, are in Mr. Rosenberg’s
possession.

There is very little difference between male and female, except
that the latter is rather smaller and not quite so bright on the
head and breast. The young bird lacks the golden hue on the
green portions of the plumage, the black of the head and throat
ismuch duller, and the beautiful golden yellow rump of the adults
is but faintly indicated by a smail patch of dull yellow.

C. johanne is another species peculiar to the humid forest
region of the Pacific coast. Besides the above specimens, and
the type (now in the Tring Museum), the only other examples
on record are two from Paramba, N.W. Ecuador, procured by
one of Mr, Rosenberg’s correspondents *.

31. CALOSPIZA RUFIGULA Bonap.

Tanagrella rufigula Bonaparte, Compt. Rend. Acad. Sci. Paris,
xxxil. (séance 20. Jan.) p. 77 (1851.—Keuador, Bourcier colli.) ;
idem, Rev. Mag. Zool. (2) iii. p. 130 (mars 1851,—reprint of orig.
descr.).

Calliste rufigularis Sclater, Cat. B. Brit. Mus. xi. p. 107 (Pasto,
Colombia t— Lehmann).

No. 2797. @ad. La Selva: 4600 ft., 15.x.09.—Wing 60;
tail 44; bill 10 mm.

“Tris dark brown, feet blue-grey, maxilla black, mandible
blue.”

This bird is identical with typical specimens from Ecuador,

La Selva is the most northerly locality for C. rufigula yet known,
though an example had already been taken by Mr. Lehmann at
Pasto in Southern Colombia, near the Ecuadorian frontier.

32. CALOSPIZA AURULENTA AURULENTA Lafr,

Tanagra (Aglaia) aurulenta Latfresnaye, Rev. Zool. vi. p. 290
(1843.—“ Colombie,” se. Bogota).

Calliste aurulenta Sclater & Salvin, P. Z. 8. 1879, p. 498 (Con-

cordia, Frontino).

* Dr. Selater’s record (Ibis, 1901, p. 597) from Peru is a mistake, cfr. Hellmayr,
Ibis, 1910, p. 328, footnote fF.
+ Not Ecuador as stated 0. cit. —

BIRDS OF WESTERN COLOMBIA. 1103

No. 3747. 9 ad. Pueblo Rico: 5200 ft., 10.ix.09.— Wing
73; tail 48; bill 10 mm.

“Tris dark brown, feet grey, bill black.”

We have also several specimens from Rio Lima, 6000 ft.,
collected by the late J. H. Batty in August 1898. They agree
perfectly with typical Bogoté skins, while birds from Western
Keuador are generally slightly paler on the head and belly.

C. aurulenta sclatert Lafr. replaces it in the Eastern Cordillera
of Colombia. It is known to me only from Bogota specimens.

33, CALOSPIZA ICTEROCEPHALA Bonap.

Calliste icterocephala Bonaparte, Compt. Rend. Ac. Sci. Paris
xxxii. (séance 20. Jan.) p. 76 (1851.—Ecuador, Bourcier coll.) ;
Sclater, Contrib. Orn. 1851, part ii. April, p. 53, pl. Ixx. fig. 1 (the
type stated to have been obtained in the valley of Punta Playa,
south of Quito); Sclater & Salvin, P. Z. S. 1879, p. 498
(Frontino).

Nos. 2825, 2837. 2 2 ad. Pueblo Rico: 2, 6.xi.09.— Wing 71,
69; tail 45, 46; bill 11 mm.

“Tris dark brown, feet grey, bill black.”

Agreeing with specimens from Ecuador and Chiriqui. C’ zetero-
cephala has a wide range, extending from Costa Rica to Western
Keuador. Jam unable to detect differences, either in size or in
colour, between skins from various localities.

34. CALOSPIZA LAVINIA LAVINIA Cass.

Calliste lavinia Cassin, Proc. Acad. N. Sci. Philad. x. p. 178
(1858.—‘ Isthmus of Darien, New Grenada ”).

C’. lavinice Cassin, Proc. Ne N. Sci. Philad. 1860, p. 142 (“ Camp
Toucey, in the mountains of the Rio Truando”—one ¢ imm.);
idem, |. ec. 1864, p. 286, pl. 1. fig. 1 (figure of type).

C. envilive Dalmast Bull. B. O. C. xi. p. 35 (1900.—San José and
El Paillon, near Buenaventura, W. Colombia); Sclater, Ibis,
1901, p. 596, pl. xii. fig. 1 (S. Javier, Rio Cachabé, N.W. Ecuador) ;
‘ofr. Hellmayr, Rev. Frang. d’Orn. no. 11, 1910, p. 161-162
(crit.).

Nos. 2149, 2159. ¢ g ad: Sipi: 2, 5.x.08.—Wing 67, 68;
tail 43, 44; bill 10 mm.

Nos. 2183, 2190. $ ¢ juv. Sipi: 12, 14.x.08—Wing 65;
tail 44; bill 94 mm.

No. 2191. Q@ ad. Sipi: 14.x.08.—Wing 63; tail 42; bill
93 mm.

“« Tris light brown, feet grey, maxilla dark, mandible lighter
brown.”

In addition, the Munich Zoological Museum possesses one
adult, one young male and two females, from San José, near
Buenaventura, 600 feet alt., which were formerly in Comte de
Dalmas’ collection and formed part of the typical series of
C. emilie Dalm. As pointed out by me in Revue Frang. @Orn.

1104 MR. C!. E. HELLMAYR ON THE

No. 11, p. 161-162, the blue-throated form is, however, the true
C. lavinia of Cassin, while the birds inhabiting Veragua and Costa
Rica will have to stand as C. l. dalmasi Hellm. C. Ll. lavinia
ranges from Darien through Western Colombia to N.W. Ecuador,
whence the Tring Museum received several specimens, obtained
by Mr. G. Flemming at San Javier, on the banks of the Rio
Cachabi. It may be mentioned that the same institution
possesses also an adult male, taken by Mr. W. F. H. Rosenberg
in 1895 at Juntas, Rio Dagua.

35. CALOSPIZA GYROLOIDES GYROLOIDES Lafr.

Aglaia gyroloides Latresnaye, Rev. Zool. x. p. 277 (1847—new
name for Aglaia peruviana (nec Tanagra peruviana Desmarest)
Swainson, Anim. in Menag. 1838, p. 356: ‘‘ Peru”—errore! we
substitute Colombia).

Calospiza gyroloides deleticia Bangs, Proc. Biol. Soc. Wash. xxt.
p- 160 (1908.—San Antonio, W. Colombia, 5800 ft.).

Calliste gyroloides Wyatt, Ibis, 1871, p. 325 (Mountain chain
between Bucaramanga and the Magdalena); Berlepsch. Journ. f.
Orn. 1884, p. 289 (Bucaramanga); Sclater & Salvin, P.Z.8.
1879, p. 499 (Concordia, 6000 ft. ; Remedios, 2360 ft.).

No. 632. ¢ ad. Jiménez, 1600 ft.; 6.vii.07.—Wing 81;
tail 55; bill 113 mm.

Nos. 2838, 3754. ¢¢ ad. Pueblo Rico, 5200 ft.; 6.xi.,
14.ix.09.— Wing 783, 82; tail 54, 57; bill 12 mm.

No. 3766. gad. Siaté, Rio Siato, 5200 ft. ; 22.1x.09.— Wing
79; tail 54; bill 12 mm.

Nos. 3753, 3767. 2 Q ad. Siaté, Pueblo Rico: 13, 22.1x.09.—
Wing 74, 75; tail 52, 51; bill 113 mm.

“ Tris dark brown, feet leaden grey, bill black.”

This series as well as other specimens from Rio Lima (Cauca
Valley, 5000 ft.), Bucaramanga and Bogota bear out the character
claimed by Mr. Bangs for his deleticia, viz., all have the lesser
wing-coverts green like the remainder of the wing. Unfortunately,
however, this form is the true gyroloides, as may be easily seen
on referring to the'original description. Lafresnaye’s name was
proposed as a mere substitute for the preoccupied term perwviana
of Swainson, who expressly says: ‘‘ shoulder-coverts green, instead
of golden yellow.” The habitat assigned to perweiana by
Swainson—Lafresnaye does not trouble himself with localities—
is, of course, erroneous since Peruvian birds, as will be shown
hereafter, possess a very large, golden yellow shoulder patch.
Swainson’s type specimen is more likely to have come from the
highlands of Colombia, to which the green-shouldered race appears.
to be confined. While I cannot, therefore, agree with Mr. Bangs
in the application of the name gyroloides, yet this author is
perfectly right in considering the Colombian and Central American
yaces as distinct. The study of the fine series in the Munich
Museum, together with other specimens lent by Count Berlepsch,
Dr. Hartert, and Dr. yon Lorenz of Vienna, shows that there

BIRDS OF WESTERN COLOMBIA. 1105

are at least three geographical forms of C. gyroloides to be dis-
tinguished. In the following lines I give a short resumé of their
Eicon which, I hope, will enable ornithologists to identify
the various subspecies.

(a) C. GYROLOIDES GYROLOIDES Lafr.

Hab. Mountains of Western and Central Colombia: Jiménez
(1600 ft.), Pueblo Rico, Siato (5200 ft.), San Antonio (5800 ft.),
province of Choco (Palmer) ; Rio Lima (5000 ft.), Cauca (Batty) ;
Concordia (6000 ft.), Remedios (2360 ft.), Antioquia (Salmon) ;
Bucaramanga (Minlos, Wyatt); Fusagasuga (6000 ft.: Wheeler) * ;
also found in Bogota collections.

Adult. Bill large and heavy; rufous chin spot immediately
followed by the cerulean blue colour of the under parts; rufous
crown without any yellow border behind or with but a very
narrow, yellowish line; lesser wpper wing-coverts green like the
median and greater ones, sometimes slightly more yellowish green
than the latter, but never yellow.

Specimens from different localities present the following
measurements :—

Four adult males from Chocé: Wing 783-82; tail 54-57 ; bill
114-12 mm.

One adult male from Rio Lima, Cauca: Wing 82; tail 54;
bill 12 mm.

One adult male from Bucaramanga (Mus. Berlepsch): Wing
78; tail 53; bill 11 mm.

Seven adults from Bogota coll.r: Wing 75-82; tail 52-56 ;
bill. 113=12 mm. *

Two females from Choco: Wing"/4, 75; tail 52, 51; bill 114,
12 mm. -

(b) C. GYROLOIDES BANGSI, subsp. n.

Hab. Costa Rica, Chiriqui, and apparently Western Ecuador.

Type in the Zoological Museum of Munich, No. 09.5340. ¢ ad.
Boquete, Chiriqui, 3500 feet, December 17, 1904. H. J. Watson
coll.
Adult. Like C. g. gyroloides. with no or very little green,
between the rufous chin-spot and the cerulean blue colour of the
under parts; yellowish border to posterior edge of rufous cap
absent or but faintly indicated : bill large and heavy ; ; but differs
in having the anterior lesser wing (or shoulder) coverts pale golden
yellow, this colour forming a conspicuous patch on the wing,
abruptly contrasting with the green of the remaining portion.

Specimens from different localities measure as follows :—

Six adult males from Chiriqui: Wing 78-80; tail 54-55;
bill 12 mm.

* Mr. Rosenberg tells me that there are two specimens of the green-shouldered
race from this locality in the British Museum, and that Salmon’s two skins from
Antioquia belong also here.

+ Four in the Munich Museum, three in the collection of Count Berlepsch.

1106 MR. C. E. HELLMAYR ON THE

One adult male from Costa Rica (Naranjo): Wing 81 ; tail 56;
bill 133 mm.

Hight adult males from Western Ecuador (Intag, 8. Nicolas,
Gualea): Wing 78-80; tail 52-58; bill 12-122 mm.

Five adult males from Paramba, N.W. Ecuador: Wing 75-79 ;
tail 52-55; bill 12 mm.

Remarks. It is a curious fact that the birds from Western
Eeuador should belong to the Central American race, and not to
C. g. gyroloides which is found in the neighbouring republic of
Colombia. However, all of the many specimens—nearly thirty
from Paramba, and eleven from other localities in Western
Kceuador—which I have been able to examine showed the pale
golden yellow shoulder-spot well-developed, thereby differing very
markedly from the Colombian form with its uniform green wings.
Although the majority of the Ecuadorian skins have the blue of
the rump rather lighter, and the top and sides of the head
somewhat clearer rufous, yet many specimens are quite indis-
tinguishable from the Chiriqui ones. I do not, of course, believe
in a discontinuous distribution, and fully expect that C. g. bangsi
will be found to exist in the Colombian coast district, having
thus an uninterrupted range from Western Ecuador to Chiriqui,
while C@. g. gyroloides is most probably confined to the high, open
country of the interior. This view is also shared by Mr. W. F. H.
Rosenberg, who has a considerable knowledge of the local distri-
bution of birds in those western districts.

The single Costa Rica male has a decidedly longer bill than any
other example examined by me.

I have named the new fog Mr. Outram Bangs, of Boston,
in recognition of his numerdUs important contributions to the
ornithology of tropical America.

(c) C. GYROLOIDES CATHARINA, subsp. n.

Hab. Upper Amazonia: from the eastern slopes of the Andes
in Colombia (Rio Meta*) and the banks of the Rio Negro
(Marabitanas, Rio Xié) through Eastern Ecuador and Peru to
Northern Bolivia (Yuracarés).

Type in the Zoological Museum of Munich, No. 11,399 ¢ ad.
Chaquimayo Carabaya, 8.E. Peru, 3000 feet, 29.viii.1910. H. &
C. Watkins coll. no. 327.

Adult. Smaller, with a much weaker, slenderer bill ; rufous cap
bordered behind by a very distinct, golden yellow band, from 2
to 3mm. wide; upper throat dark green, separating the rufous
chin-spot from the blue foreneck ; the whole of the lesser and the
adjoiming median upper wing-coverts deep orange golden, forming
a very large, bright shoulder-patch. ‘'I'his patch is about twice as
large asin OC. g. bangsi, and of a much deeper, more orange golden
tinge.

* According to Mr. Rosenberg, the British Museum has four adult birds of this

form from the Rio Meta (800-1500 ft.; Wheeler coll.) and several others obtained
by Buckley in Eastern Ecuador.

BIRDS OF WESTERN COLOMBIA. 1107

Specimens from different localities measure as follows :—

Three adult males from Chanchamayo (C. Peru): Wing 75-77;
tail 49-53 ; bill 10-11 mm.

One adult male from N. Peru (Huayabamba).—Wing 73;
tail 48; bill 11 mm.

Four adult males from 8.E. Peru (Marcapata): Wing 73-76 ;
tail 52-54; bill 11 mm.

Three adult males from Upper Rio Negro*: Wing 70-72: tail
47-493; bill 10 mm.

Two adult males from the Rio Putumayo, §.E. Colombia
(Mus. Berlepsch): Wing 703, 71: tail 49, 50; bill 10-104 mm.

Two adult males from Bogota collections: Wing 75, 76; tail
53, 54; bill 103, 11 mm.

Remarks.—These fifteen examples differ very conspicuously
from C. g. bangsi in the characters given above. The three from
the upper Rio Negro and one from the Putumayo have the head
of a deep chestnut-brown (instead of cinnamon-rufous) colour, but
the second specimen from the Rio Putumayo and the two Bogota
skins agree with the Peruvian ones, The dimensions are also
rather variable, though apparently not connected with any
particular geographical area.

I have named this well-characterized new form after my wife,
who takes considerable interest in ornithology and has materially
helped me in the preparation of this report.

36. CALOSPIZA PALMERI Hellm.

Calospiza palmeri Hellmayr, Rev. Frang. dOrnith. No. 4, p. 49
(1909.—Sipi, W. Colombia); idem, Ibis, 1910, p. 330, pl. v.

Nos. 2164, 21667, 2167, 2186. ¢ ¢ ad. Sipi, Rio Sipi:
7, 13.x.08.—Wing 803-83 ; tai] 554-60; bill 10-11 mm.

No, 2165. 9 ad. Sipi: 7.x.08.—Wing 763; tail 50; bill
11 mm.

‘“‘ Tris dark brown, feet and bill black.”

This fine new bird was fully described by me J. c. and is faith-
fully depicted on plate v. in ‘ The Ibis,’ 1910.

Mr. Palmer has not succeeded in getting additional specimens
of this Tanager.

37. CALOSPIZA LARVATA FANNY Lafr.

Aglaia Fanny Lafresnaye, Rev. Zool. x. p. 72 (1847—* in Nova
Grenada ” (Delattre)—sc. Buenaventura, W. Colombia.)

Calliste francesce Cassin, Proc. Acad. N. Sci. Philad. 1860,
p. 142 (Turbo, N.W. Colombia).

* Two from the Rio Xié, June 1831, one from Marabitanas, March 1831, obtained
by J. rine Vienna Museum [= Calliste gyroloides Pelzeln, Orn. Bras. iii. 1869,

. 207].
P + Type of species.

{ The type is in the Museum of the Academy of Natural Sciences in Phiiadelphia
(cf. Stone, Proc. Acad. N. Sci. Philad. v. 1899, p.51),and not in Baron de Lafresnaye’s
Collection, as erroneously stated by Ridgway (Bull. U. S. Mus. no. 50, pt. ii. p. 50).

1108 MR. C. E. HELLMAYR ON THE

O. larvata francesce Hartert, Nov. Zool. v. 1898, p. 482
(Cachabi, N.W. Ecuador).

Calliste larvata (errore, nec Dubus) Sclater & Salvin, P. Z.8.
1879, p. 499 (Remedios, Antioquia).

Nos. 2050, 2062. ¢ gd ad. Noanama: 2, 4.ix.08.— Wing 69 ;
tail 474-49 ; bill 10 mm.

Nos. © 2063, 2199, 22005 9 Q ad? (iNoanamal: ) Arxe a aite:
19.x.08.—Wing 65-682 ; tail 44-47 ; bill 10 mm.

No. 2408. 2 ad. Ndévita: 17.xii.08.—Wing 653; tail 46;
bill 11 mm.

No. 2032. $ juv. Noanama: 28.vi1i.08.

‘“‘ Tris dark brown, feet and bill black.”

This is the true C. J. fanny which was originally described from
specimens obtained by Delattre in the same district of Western
Colombia. The comparison of these birds with the large
series in the Munich Museum clearly proves that the Central-
American form has been wrongly referred to C. l. fanny by
Sclater, Ridgway, and others. (. 1. fanny as represented by
specimens from Western Colombia and N.W. Ecuador (which
are exactly similar) is an exceedingly well characterized race. The
greater upper wing-coverts and the remiges are either uniform
black or show only the faintest trace of greenish fringes in their
apical half; the lower back and rump are very pale sky-blue ; the
blue frontal patch is very strongly developed, reaching as far as
the posterior edge of the eye; the flanks are washed with pale
sky-blue, without any purple-blue admixture; the bill is small
and slender. Birds from Chiriqui and 8.W. Costa Rica (Guana-
caste, Pozo Azul de Pirris), however, differ at a glance in having
the greater wing-coverts and remiges very distinctly margined
with pale green, and the flanks much more extensively and darker
blue, with a strong purplish blue tingeanteriorly. Moreover, the
blue frontal patch is much more restricted, never reaching further
back than to above the middle of the eye, and the bill is larger
and stouter. In all of these points they agree with C. 1. larvata
Dubus*, of which we have a fair series from Guatemala and
Eastern Costa Rica (Carrillo), but may easily be distinguished
from that form by their coppery-golden, instead of deep coppery
reddish-brown throat, much paler blue cheeks with very little
purplish tinge, and slightly paler blue rump.

CO. 1. fanny is restricted to the forest-belt bordering the Pacific
coast, ranging from N.W. Ecuador (Cachabi, 8. Javier, Pambilar)
to Turbo, on the Gulf of Uraba, N. Colombia.

38. CALOSPIZA RUFICERVIX RUFICERVIX Prév. & Des Murs.

Tanagra ruficervix Prévost & Des Murs, Voyage de la Vénus,
Atlas, Oiseaux, pl. v. fig. 1 (1846—no locality).
T. (Calliste) rufivertex iidem, 1. c., Zool. (text), V, 1, p. 212

* Calliste larvata Dubus, Esquiss. Ornith. pl. 9 (1845(?).—Tabasco, $.K. Mexico).

BIRDS OF WESTERN COLOMBTA. 1109

(1855. —The type stated to be from Guatemala, which is doubtless
a mistake),

Calliste ruficervix Sclater & Salvin, P. Z. 8. 1879, p. 499 (Con-
cordia).

Nos. 2813, 2824. 9 2 ad. Pueblo Rico: 20, 26.x.02.—Wing
72, 70; tail 48; bill 10 mm.

‘“‘ Tris dark brown, feet grey, bill black.”

The Munich Zoological Museum also possesses an adult male
from Rio Lima, 5000 ft., taken in August 1898 by the late
J. H. Batty.

The birds from Western Colombia are in every respect similar
to others from Bogota and Western Heuador in our collection.*

Cr. ruficervix is found only in Colombia (Western Cordillera
and Bogota-coll.) and Western Ecuador (Pallatanga, Cayandeled),
living at rather high altitudes. Im Hastern Hcuador (Machay,
Napo) it is represented by the well-characterized C. r. taylort
Tacz. & Berl. t, while, in Peru and Northern Bolivia, C. r. fulvi-
cervic Scl. & Salv.{ takes its place. Of this last-named
form, the Munich Museum possesses specimens obtained by
Mons. G. A. Baer at Nuevo Loreto, Northern Peru, in June
1900.

39. CALOSPIZA LABRADORIDES Boiss,

Tanagra (Aglaia) labrudorides Boissonneau, Rev. Zool. iti. p, 67
(1840.—Santa-Fe-de- Bogota).

Calliste labradorides Sclater & Salvin, P. Z.8. 1879, p. 499 (Con-
cordia, Santa Elena).

No. 3751. 9 ad. Pueblo Rico: 9.ix.09.— Wing 65; tail 47 ;
bill — mm.

‘“‘ Tris dark brown, feet grey, bill black,”

We have also an adult male procured by J. H. Batty at Rio
Lima, 5000 ft., in August 1898. The birds from the Western
Cordillera agree perfectly with a series of Bogota skins.

C. labradorides is peculiar to the mountains of Colombia.

40. BuTHRAUPIS MELANOCHLAMYS Hellm.

B. melanochlamys Hellmayr, Bull. B. O. C. xxv. no. elxi.
p- 112 (June 1910.— La Selva, W. Colombia),

No. 2796. Q ad. La Selva, Rio Jamaraya, 4600 ft., 15.x.09.
Type of species. No. 10.2378, Coll. Zoological Museum, Munich.
Head all round, back and scapulars deep black, with a faint
silky gloss; lower rump and upper tail-coverts rather dull indigo-

* None of the many specimens I have examined showed any trace of the white
auricular spot or any white on the lesser wing-coverts, characters given by
Dr. Sclater tor his Calliste lewcotis (Contrib. to Ornith. 1851, pt. ii. April, p. 58.—
Kenador).

+ Calliste taylori Taczanowski & Berlepsch, P. Z.S. 1885, p. 75 (1885.—-Machay,
E. Ecuador).

* Calliste fulvicervix Sclater & Salvin, P.Z.S, 1876, p. 354, pl. xxx. fig. 1
(Aug. 1876.—Tilotilo, N. Bolivia.)

Proc, Zoot, Soc.—1911, No. LXAXY. (6)

1110 MR. C. E. HELLMAYR ON THE

blue (Valette’s Code des Couleurs, no, 417); lesser and median
upper wing-coverts uniform bright indigo-blue (im shade between
nos. 411 and 412 of Valette’s Code), forming a large pale blue
shoulder-patch ; greater series dusky, on the outer web washed
with dull bluish; primary coverts, remiges and rectrices blackish,
narrowly fringed with dull bluish along outer margin. Below:
throat and sides of breast deep black, like the back; flanks duller
and inclining to blackish slate-colour; rest of ander parts bright
saffron-yellow, passing into a deeper, more orange tinge on the
middle of the chest. Axillaries pale yellow, under wing-coverts
yellowish white.

Bill uniform black. ‘“ Iris brown, feet black.”

Wing 90; tail 53; bill 154 mm.

Mr. Palmer obtained a single specimen of this distinct species
on the San Juan slopes of the Western Cordillera. It belongs to
that section embracing B. arcei Scl. & Salv.* and B. rothschildi
Berl.t, but differs conspicuously in coloration. The most im-
portant characters of B. melanochlamys are the bright, pale blue
shoulder-patch, altogether absent in both of its allies, and the
deep black colour of the head, mantle, and scapulars, without any
bluish tinge. B. arcwi resembles it in having the breast and belly
bright yellow, but the sides and flanks, instead of being exten-
sively uniform deep black, are merely clouded with dusky, and
the upper parts as well as the throat are strongly glossed with
dark purplish blue. 3B. rothschildi, from the, foot-hills of North-
Western Ecuador, has, like the new species, the head and throat
deep black; however, the back, scapulars, upper wing- and tail-
coverts are conspicuously glossed with purplish blue, while the
under parts, with the exception of an orange-yellow patch on the
foreneck, and the lemon-yellow anal region and under tail-coverts,
are black, with a more or less distinct purple-blue sheen.

The bill is deep black in all three species which eventually
will prove to be geographical representatives. Their range is
singularly restricted :—

(a) B. arcei Sel. & Salv. inhabits the Cordillera del Chucu,
Veragua.

(b) B. melanochlamys Hellm. is found in the Western Cordillera
of Colombia, on the sources of the San Juan River.

(c) B. rothschildi Berl. is met with in the foot-hills of North-
Western Ecuador: Cachabi (450 ft.), Rio Pichiyacu (500 ft.), ete.

41. BuTHRAUPIS AUREOCINCTA Hellm.

B. aureocincta Hellmayr, Bull. B.O.C. xxv. no. elxi. p. 111
(June 1910.—Tatama Mountain, W. Colombia).
No. 3487. gad. Tatama Mountain, Chocé, 6700 feet, 8.x.09.
Type of species. No. 10.2377. Coll. Zoological Museum, Munich.
3 ad. Top and sides of the head, and nape deep black, with the
* P.Z.S. 1869, p. 439, pl. xxxi. (Cordillera del Chuct, Veragua).

+ Bull. B. O.C. vii. p. ui (Oct. 1897.—Cachabi, N.W. Ecuador) ; Hartert, Nov.
Zool. v. 1898, p. 482, pl. u. fig. 2.

BIRDS OF WESTERN COLOMBIA. WL

exception of a broad, bright saffron-yellow band which starts from
above the eye and descends the sides of the neck where it joins
the yellow malar stripe. Back bright olive-green ; upper wing-
coverts dull indigo-blue, those of the greater series dusky on the
inner web; primary coverts and remiges black, edged with dull
indigo-blue, the innermost secondaries washed with the same
colour on both webs; rectrices blackish, exteriorly broadly mav-
gined with bright olive-green. Cheeks and malar region bright
saffron-yellow ; large chin-spot pale yellow, some of the feathers
with slight dusky edges; throat and sides of the foreneck black,
the feathers of the former with half-concealed, subterminal spots
of pale yellow; a large, bright golden-yellow patch in the middle
of the foreneck; remainder of under parts olive-green, lighter
and more yellowish than the back; under tail-coverts lemon-
yellow, with the basal half olive-green. Avxillaries dusky, tipped
with greenish ; under wing-coverts greyish white.

“Tris brown, feet dark brown, maxilla black, mandible yellow.”

Wing 93; tail 60; bill 14 mm.

This species, of which Mr. Palmer sent only a single adult bird, is
most nearly related to B. edwardsi Elliot.* The two species agree
in the colour of the back, wings and tail, in having the lbpanets and
abdomen olive-green, with a large, golden- yellow patch on the
foreneck, also in rhe shape and coloration of the bill; but
BL. edwardsi may be distinguished at a glance by having the sides
of the head uniform pale blue, without any yellow or black.

B. aureocincta is known only from the Tatama Mountain, on
the sources of the San Juan River, in the Western Cordillera of
Colombia.

B. edwardsi is an inhabitant of the high mountains of Southern
Colombia and Northern Ecuador. The type, in the Paris Museum.
was brought by M. Triana from an uncertain locality in ‘‘ New
Grenada.” A single specimen was obtained by Prof, Orton at
Chillo, on the western slope of the voleano Antisana, 10,000 ft.
alt.7, Ecuador, and the British Museum received three examples
from Pasto, 8. Colombia ¢, through Mr. Lehmann. Count Ber-
Jepsch possesses _a female picked out from a lot of Quito-skins.
There is, in the United States Museum at Washington +, a skin
said to be from Esmeraldas, N.W. Ecuador, but this locality is,
no doubt, incorrect.

42, CoMPSOCOMA NOTABILIS Jard.

Tanagara (sic) notabilis Jardine, New Edinburgh Philos. Journ.
(n. s.) 1. p. 119 (July 1855.—KHastern Cordillera, Ecuador.
W. Jameson coll.).

Nos. 3785, 3786. $$ ad. Tatamda Mountain, 6700 ft.,
8.x.09.— Wing 99, 100; tail 82, 83; bill 19 mm.

«Tris brown, feet black, maxilla black, mandible blue.”

* Nouv. Arch. Mus. Paris, i. Bull. p. 77, pl. iv. fig. 2 (1865.—“ Nouvelle Grenade , ).

t Salvin, Ibis, 1874, p. 307.

{£ Not Ecuador, as stated in the Cat. B. Brit. Mus. xi. p. 150.
oe

WW MR. C. E. HELLMAYR ON THE

The specimens correspond to the original description, and to
the figure in P. Z.S. 1855, plate xc.

So far as I know, this beautiful species has not been recorded
before from Colombia, and was reported only as an inhabitant of
the highlands of Northern Ecuador.

43, TANAGRA PALMARUM MELANOPTERA Scl,

[Tanagra palmarum Wied, Reise nach Brasil. 11. p. 76 (1821.—
Canavieras, Bahia. |}

T. melanoptera (Hartlaub MS.) Sclater, P. Z. 8. 1856, p. 235
(Jan. 1857.—Kastern Peru, etc.).

T. palmarum (nec Wied) Sclater & Salvin, P.Z.S5. 1879,
p- 500 (Remedios).

T. p. melanoptera Berlepsch, Journ. f. Orn. 1884, p. 291
(Bucaramanga).

Nos. 2499, 2650, 2682. g gad. Ndévita: 281.09; Tadeo:
18.v., 1.vi.09.—Wing 92; tail 70-72; bill 13 mm.

No. 2805. g ad. Loma Hermosa, 4180 ft., 23.x.09.—Wing
94; tail 73; bill 13 mm.

Nos. 2424, 2649, 2716. Qad., Qjr., ¢ juv. Névita: 22.xi1.08;
Tad6: 18.v., 18.vi.09.—Wing 89-90; tail 72, 70, 67; bill 123—
13 mm.

“Tris dark brown, feet grey, bill black.”

The adult birds agree with topotypical skins from Northern
Peru and others from Eastern Ecuador (Napo), Venezuela, Trini-
dad, ete., except that the top of the head is of a paler, duller
yellowish olive. In this respect they resemble a series of Bogota
skins. It must be mentioned, however, that two or three from
Bogota have the eap quite as bright olive-yellow as the Peruvian
ones. More information about the geographical distribution of
these forms is required before any further separation can be
attempted.

T. p. violilavata Berl. & Tacz.,* from Western Ecuador, is very
different from the Colombian birds. It has both the upper and
lower parts much more strongly suffused with violet, and lacks
the olive-yellowish cap, the pileum being bluish like the back.

44, RAMPHOCELUS ICTERONOTUS Bonap.

Ramphocelus icteronotus Bonaparte, P. Z.8, 1837, p.121 (1838,
June.—loc. ign., type in Paris Museum); Cassin, Proc. Acad.
N. Sci. Philad. 1860, p. 141 (Turbo, Rs. Atrato and Truando).

Rhamphocelus icteronotus Sclater & Salvin, P. Z.S. 1879, p. 501
(Remedios, Neche).

Nos. 1969, 2207, 2407. § g ad. Guineo: 5.viii.07; Novita:
17.x11.08; Noanama: 21.x.08.— Wing 85-87 ; tail 72-78; bill 15—
16 mm.

No. 2433. § vixad. Novita: 24.xii.08.—Wing 85; tail 75;
bill 15 mm.

* P.Z.S. 1883, ». 546 (1883.—Chimbo, 8.W. Ecuador).

BIRDS OF WESTERN COLOMBIA. AS

No. 2110. g imm. Sipi: 23.1x.08— Wing 85; tail 77;
bill 16 mm.

Nos. 1956, 2228, 2240. 9 9. San Joaquim, 1.viii.08; Rio
Cajon: 2,5.xi. 08.—Wing 79-82; tail 71-76; bill 15-16 mm.

“Tris red, dark brown (no. 2433), feet blue-grey or blue,
bill blue.”

The Munich Museum also possesses specimens obtained by the
late J. H. Batty on the Rio Guapi, 200-300 feet alt.

R. icteronotus ranges from Veragua to S.W. Ecuador (district
of Guayaquil) *, frequenting the forest-region, from sea-level up
to about 3000 feet.

45, PIRANGA RUBRA RUBRA Linn.

Fringilla rubra Linneus, Syst. Nat. x. p. 181 (1758,— based
on Catesby, Nat. Hist. Carolina ete. i. p. 56, pl. 56; Carolina,
Virginia).

Pyranga estiva Cassin, Proe. Acad. N. Sci. Philad. 1860, p. 140
(Turbo); Sclater & Salvin, P.Z.S. 1879, p. 502 (Concordia,
Frontino).

Nos. 2203, 2213. ¢ g¢ ad. Noanama, 100 ft., 20, 22.x.08.—
Wing 94, 95; tail 74, 76 mm.

No. 2223. gad. Rio Cajon : 30.x.08.—Wing 96; tail 73 mm.

Nos. 2817, 2823, 2830. g¢ gad. Pueblo Rico, 5200 ft., 28, 30.x.,
3.x1.09.— Wing 93, 96, 99; tail 72, 74, 76 mm.

No. 2204. g juv. Noanama: 20.x.08.

Nos.2224, 2251, 2390. 9 9. Novita: 9,11.xii.08; Rio Cajon:
30.x.08.— Wing 92-93 mim.

“ Tris dark brown, feet grey or purplish grey, maxilla yellowish
brown or horn-colour, mandible yellowish brown, yellow, or grey.”

A common winter visitor to South America.

46. PH@NICOTHRAUPIS CRISTATA Lawr.

Phenicothraupis cristata Lawrence, Ann. Lye. N. H. N. Y. x1.
‘p. 70 (Feb. 1875.— Bogota).

Phenicothraupis cristata Sclater & Salvin, P. Z.S. 1879, p. 50
(Frontino, Antioquia); Bangs, Proc. Biol. Soc. Wash. KERSTIN 0
(Naranjito, R. Dagua).

Nos. 3778, —. 6 dad. La Selva, 4600 ft., Oct. ?09.— Wing
97, 98; tail 85, 89; bill 18, 19 mm.

No. — (not numbered). dg ad. Pueblo Rico, 5200 ft.,
Noy. 09.—Wing 98; tail 89: bill 173 mm.

No. — (not numbered). g ad. Loma Hermosa, 4150 ft.,
Oct. 20, 09.—Wing 100; tail 90; bill (damaged) mm.

Nos. 2739, 2829. 9 9 ad. Pueblo Rico: viii, 3.x1.09.—Wing
94,95; tail 86; bill 18, 20 mm.

2
5

* Ridgway (Bull. U.S. Mus. no. 50, pt. ii. p. 114) erroneously included “ Central
Peru” in the habitat of this species. Palmal whence Taczanowski (P.Z.S. 1877,
p. 332) recorded a couple, is not in Peru, but in the district of Santa Rosa, province
of Guayaquil, S.W. Ecuador. . icteronotus does not occur anywhere in Peru.

1114 MR. C. E. HELLMAYR ON THE

No. 3752. 2 juv. Pueblo Rico: 13.ix.09.—Wing 88; tail 81;
bill 17 mm.

‘Tris brown (adult), grey (juv.), feet brown, bill black.”

The adult males have the back deep vinous red, the upper
wing-coverts and outer webs of the quills scarcely duller. The
females are smaller; the crest is duller and more of a scarlet red ;
the throat rather lighter red; the back and wings are much
duller, brownish vinaceous ; breast and abdomen decidedly paler
pink, with the greyish bases of the feathers more plainly showing
through, etc.

The young bird has the upper parts dusky brown, with the tail-
coverts and rectrices dull testaceous, the lower surface dingy
cinnamomeous passing into dark brown on throat and foreneck.
There is no trace of the red crest, only a few of the crown-feathers
are narrowly edged with dull reddish.

P. cristata is peculiar to the mountainous districts of Western
Colombia. It is only occasionally met with in Bogota collections.

47. CHLOROTHRAUPIS OLIVACEA Cass.

Orthogonys olivaceus Cassin, Proc. Acad. N. Sci. Philad. 1860,
p- 140 (1860.—* Cordilleras Mountains, on the River Truando,
New Grenada”); Sclater & Salvin, P.Z.S. 1879, p. 502 (Reme-
dios, Neche).

Chlorothraupis olivacea Hartert, Nov. Zool. v. 1898, p. 483
(Cachabi, N.W. Ecuador).

Nos. 2137, 2232: $ g ad. Sipi: 11-x.; Rio Cajon: 2.x1.08.—
Wing 92, 94; tail 633, 65; bill 183, 193 mm.

No. 2379. ¢g imvm. Novita: 9.211.08—Wing 89; tail 63;
bill 183 mm.

Nos. 1962, 2101, 2111, 2123. 9 Gad. & imm. 8. Joaquim:
3.vili. ; Sipi: 21, 23, 26.1x.08.—Wing 85, 86, 86, 90; tail 59-60,
bill 18-20 mm.

“Tris dark brown, feet blue or dark grey, maxilla black,
mandible blue.”

Ndult males are dark brownish green above and have the
feathers of the pileum broadly centred with blackish; the lower
parts are nearly uniform olive-green except the upper throat,
which is light yellow. The females are much smaller, lighter
green above, and more yellowish green below with the whole
throat and middle of foreneck pale yellow; the pileum is uniform
olive-green like the back.

C. olivacea is peculiar to the humid forest-district of Western
Colombia and N.W. Ecuador.

48, HerprosPincus xAnTHopyctus Sel,

Tachyphonus wanthopygius Sclater, P. Z. 8. 1854, p. 158,
pl. lxix. (Apr. 1855—“in Nova Grenada”; descr. 2); idem, le.
1855, pl. xc. (deser. ¢ ; Bogota) ; Cassin, Proc. Acad, N. Sci. Philad.

BIRDS OF WESTERN COLOMBIA. WW

1860, p. 142 (Truando; one ¢ adult); Sclater & Salvin, P. Z. 8.
1879, p. 503 (Remedios).

Nos. 2357, 2709. $ gad. Novita: 1.xii.08; Tado: 16.v1.09.—
Wing 92, 89; tail 68, 65; bill 18 mm.

Nos. 2253, 2358, 2610. 9 @ ad. Novita: 9.xi., 1.11.08;
Condoto : 17.iv.09.—Wing 88-90; tail 65-67; bill 16-18 mm.

“Tris dark red, feet dark grey, bill black.”

These birds are identical with Bogotd-skins in the collection of
the Munich Museum. Like the latter, the males have a broad
scarlet supra-auricular stripe, and the lesser upper wing-coverts
bright lemon-colour,

H. wanthopygius ranges from N.W. Colombia (Truando) south-
wards to S.W. Ecuador, where a specimen was obtained by
Dr. Siemiradzki in the vicinity of Chimbo*. In Panama and
Eastern Costa Rica it is represented by the nearly allied, and
perhaps only subspecifically distinct H. rubrifrons Lawy.t
Although considered by Salvin¢ to be inseparable from //. xan-
thopygius of Colombia, two birds from Veragua and Chiriqui
are so much smaller (wing 78, 80; tail 58 mm.) than any of
the numerous (about fifteen) specimens of the southern form, that
there can be scarcely any doubt as to their distinctness. More-
over, in H. rubrifrons the sexes are alike, the male lacking the
scarlet supra-auricular stripe as well as the yellow patch on the
lesser wing-coverts. Cfr. Ridgway, Bull. U.S. Nat. Mus. no. 50,
pt. 11. pp. 104-105.

49, TACHYPHONUS DELATRII Lafr.
Tachyphonus Delatrit Lafresnaye, Rev. Zool. x. p. 72 (1847.—

Buenaventura, W. Colombia).

T. De Lattrei Cassin, Proc. Acad. N. Sci. Philad. 1860, p. 142
(Falls of the Truando).

T. delattrii Sclater & Salvin, P. Z.S. 1879, p. 503 (Remedios,
Neche).

No. 1959. g ad. San Joaquim (Buenaventura): 1.viii.08.—
Wing 74; tail 64; bill 14 mm.

Nos. 2027, 2064. ¢ gad. Noanama: 28.viii., 4.ix.08.— Wing
73, (2; tail 65, 63; bill 13, 14 mm.

INos. 2222, 2238, 3 oad) Rio Cajon: 30.x., 3.x1.08.— Wing
TA. (38> tall Gos G3 sbullil5. 14s mmo:

Nos. 2094, 2109, 2117, 2139.9 G ad. Sipi: 19, 23, 241x.;
1.x.08.—Wing 71-73; tail 62-64, bill 13-14 mm.

Nos 2090M 2180.) 2Gsam eae ClO 8 sip 18.) BOs ails
13.x.08.— Wing 63-67; tail 55-58; bill 12-13 mm.

“Tris dark red, feet and bill black.”

These examples are topotypical, Lafresnaye’s type having been

* Berlepsch & Taczanowski, P. Z.S. 1883, p. 547: Tachyphonus xanthopygius.

+ Tachyphonus rubrifrons Lawrence, Proc. Acad. N. Sci. Philad. xvii. p. 106
(1865.—Line of Panama Railroad).

+ Ibis, 1870, pp. 109-110: 7. propinguus.

1116 MR. C. EK. HELLMAYR ON THE

secured by Delattre at Buenaventura, in the Bay of Choco.
Birds from Western Ecuador are absolutely similar.

Skins from Costa Rica (Carrillo etc.) are larger, with a longer
and more slender bill; the males are of a duller, more brownish
black, and have the vertical crest somewhat deeper orange.
They are perhaps subspecifically separable. The locality ‘ Pasto’ *
is, no doubt, incorrect, for 7. delatrii is found only in the
lowlands and foot-hills.

50. Mirrosprneus cassinit Lawr.

Tachyphonus Cassinti Lawrence, Ann. Lye. N. H. N.Y. vu.
p- 297 (Jan. 1861.—Panama, Railroad).

Tachyphonus ¢ — Cassin, Proc. Acad. N. Sci. Philad. 1860,
p. 142 (Falls of the Rio Truando).

Eucometis cassini Sclater & Salvin, P.Z.S8. 1879, p. 503 (Neche);
Hartert, Nov. Zool. v. 1898, p. 483 (Cachabi, N.W. Ecuador).

Nos. 2612, 2613. g g ad. Condoto: 20.iv.07.— Wing 86, 84;
tail 77, 74; bill 18 mm.

“Tris grey, feet black, bill black, mandible grey.”

These birds agree, in coloration and size, with others from
Costa Rica. Topotypical Panama specimens are not available for
comparison.

M. cassinii ranges from Costa Rica through Western Colombia
southwards to N.W. Ecuador, whence Mr. Rosenberg’s collectors
have sent many specimens*. It is an inhabitant of the humid
forests of the lowlands and foot-hills, up to about 2000 feet.

51. Hem1rHRaAvPis SALMONI Scl.

Dacnis salmoni Sclater, Cat. B. Brit. Mus. xi. p. 27, pl. i.
fig. 2 (1886.—Remedios, Antioquia; = Q); efr. Hellmayr, Nov.
Zool. xiii. 1906, p. 317 (crit.).

Nemosia rosenbergi Rothschild, Bull. B. O. C. vii. p. vi (Oct.
1897.—Cachabi, N.W. Ecuador; = dad.); Hartert, Nov. Zool.
v. 1898, p. 483, pl. 1. fig. 1.

Not DRG OPUS) BOL WR i gf acl Iowan 9) 75 Zlleaxi..
11.xi1.01.— Wing 6 REA 76 ¢ tail 50-53 ; bill 123-123 mm.

No. 2096. ¢ ad. Near ‘Sipi : 19.ix.08.— Wing 68; tail 50 mm.

No. 2261. ¢ vix ad. Novita: 11.x1.08.—Wing 66; tail 49;
bill 185 mm.

Nos. 2324, 2331. ¢ ¢ juv. Novita: 21, 24.x1.08— Wing 65;
tail 49, 50; bill 123, 133 mm.

Nos. 2323, 2337. 9 9 ad. Novita: 21, 25.x1.08 —Wing 957;
tail 42, 43; bill 11-12 mm.

“ Tris dark brown, feet dark brown (d ad.), grey (dg juv. and 2 );
maxilla black, rmaenible light brown or yellowish pense ad.),
grey (do juv. and Qe

* Cat. B. xi. p. 215.

+ Ridgway (Bull. U.S. Mus. no. 50, pt. ii. p. 169) mentions, as existing in tie

U.S. National Museum, a female from“ Guayaquil, S.W. Eenador, but this locality
requires confirmation.

BIRDS OF WESTERN COLOMBIA. iplele7s

[Mus. Brit. 2 imm. Remedios, Antioquia. Type of D. salmon.
—Wing 58;, tail 42; bill 12 mm.]

The splendid series forwarded by Mz. Palmer leaves no longer
any doubt that Dacnis salmoni is the female of WV. rosenbergi.
The females from Novita agree perfectly with the type in the
British Museum, with which they were kindly compared by my
friend Dr. Hartert. Moreover, they correspond exactly to a
detailed description drawn up by me when in London some years
ago. The type of WV. rosenbergi was obtained at Cachabi, N.W.
Keuador, 450 feet, in November 1896, by Mr. W. F. H. Rosenberg,
and is faithfully depicted on the plate quoted above.

The female may be described as follows :—Above dull yellowish
olive, pileum rather darker, hind crown somewhat shaded with
cinereous; upper wing-coverts like the back; quills dusky,
exteriorly edged with yellowish olive ; rectrices dusky, washed
with pale yellowish olive; sides of the head dull olive; throat
and foreneck dingy buff; remainder of under parts bufty white,
niore purely white along the middle of the abdomen ; under tail-
coverts light buff; axillaries and under wing-coverts white ;
distinct margin along the inner web of the remiges olive-yellow.
Wi ings and tail are much shorter than in the adult male, and the
bill is hkewise shorter, with the lower mandible pale hoa brown
instead of whitish.

The young males (nos. 2324, 2331) in coloration resemble the
female, but are very nearly as large as the adult males. They
show a few orange-yellow feathers on the sides of the head. The
mandible is pale brown as in the females.

In structure, H. salmoni agrees with H. chrysomelas Scl. &
Salv. *, from Southern Central America. The females also
present the same style of coloration, that of H. salmoni being,
however, easily recognizable by having the throat and foreneck
buff (instead of olive- -yellow), the belly buffy white (not pale
yellowish), and the upper parts of a considerably darker tinge.

While H. salmon and H. chrysomelas ave strictly congeneric,
T am a little doubtful whether they can be kept in the genus
Tlemithraupis, for the typical species, H. ruficapilla Vieill., and
its allies have a much shorter, ibaendler, less constricted bill.

H. salmoni is peculiar to the humid forest-districts of Western
Colombia and North-western Ecuador.

52. BUARREMON BRUNNEINUCHA Lafr.

Embernagra brunneinucha Lafresnaye, Rev. Zool. 11. p. 97
(1839.—‘“ Mexico”).

Buarremon brunneinuchus Sclater & Salvin, P.Z.S. 1879,
p- 504 (Concordia, Medellin, Santa Elena).

No. 3743. gad. Pueblo Rico: 9.1x.08.—Wing 81; tail 83;
bill 173 mm.

* Tachyphonus chrysomelas Sclater & Salyin, P. Z.S. 1869, p. 440, pl. xxxii.
(Cordillera del Chuet, Veragua)..

1118 MR. C. E. HELLMAYR ON THE

“Tris and feet dark brown, bill black.”
Identical with specimens from Bogota and Venezuela (Cumbre
de Valencia, Mérida). Typical Mexican birds are not available.

Cfr. Ridgway, Bull. U.S. Mus. no. 50, pt. i. p. 466.

53. ARREMON AURANTIIROSTRIS OCCIDENTALIS, subsp. n.

No. 2557. S$ vixad. Juntas, Rio Tamana (405 ft.), 26.11.09.—
Wing 73; tail 56; bill 153 mm.

No. 2573. gad. Condoto, R. Condoto (150 ft.), 27.11.09.-—
Wing 73; tail 58; bill 144 mm.

‘“‘ Tris dark brown, feet pink, bill bright orange-red.”

Messrs. Salvadori and Festa * have most correctly pointed out
the differences between specimens from Western and Eastern
Eeuador. The large series in the Munich Museum, however,
proves that the birds from the western slopes of the Andes are
clearly distinct from A. erythrorhynchus of Bogota, and constitute
a third, hitherto overlooked race. <A. spectabilis, A. erythro-
rhynchus, and the new form are evidently the southern repre-
sentatives of the Central American A. auraniiirostris Lafr. 7 and
should, according to my view, be ranked as subspecies of the
latter. The northern bird may, however, readily be distinguished
by its much larger, heavier bill, much broader, black jugular band,
and pure olive-green black.

Characters, range and synonymy of the three races are as
follows :—

(a) A. AURANTITROSTRIS SPECTABILIS Scl.

Arremon spectabilis Sclater, P. Z. 8. 1854, p. 114, pl. Ixvi.
(Apr. 1855.—Quijos, Hast Ecuador); idem, P. Z. 5. 1856, p. 82
(Quixos); idem, P. Z. 8S. 1858, p. 72 (Rio Napo, E. Ecuador) ;
idem, Cat. B. Brit. Mus. x1. 1886, p. 275 (part.: e-g, Rio Napo;
h, Sarayacu; p, Eastern Peru); Salvadori & Festa, Boll. Mus. Zool.
Torino, xiv. no. 857, 1899, p. 20 (Rio Santiago and R. Zamora,
E. Ecuador).

A. erythrorhynchus (nec Sclater) Berlepsch & Taczanowski,
P. Z.S. 1883, p. 548 (Huambo, E. Peru; in text of A. spectabilis
no. 49); Taczanowski, P. Z.S8. 1882, p. 16 (Huambo); idem,
Orn. Pérou, ii. 1884, p. 535 (Huambo).

Habitat. Easrern Ecuapor: Quijos (fide Gould), Rio Napo
(fide Verreaux), La Concepcion (Petit), Sarayacu (Buckley),
R. Santiago and R. Zamora (esta); Norra Peru: Huambo
(Stolzmann).

Characters. Back, larger upper wing-coverts, and outer webs of
remiges deep brownish or rufescent olive; bend of the wing
bright orange; bill small and stout. Wing 70-73; tail 58-60;
bill 123-133 mm.

Material. A series from La Concepcion, Rio Napo, East
Keuador, obtained by Louis Petit, in the Munich Museum.

* Boll. Mus. Zool. Torino, xiv. no. 357, 1899, p. 21.
+ Rey. Zool. x. p. 72 (1847.—Panama).

BIRDS OF WESLERN COLOMBIA. 1119

Observations. As already stated by Dr. Sclater and more
recently by Salvadori and Festa, birds from Eastern Ecuador
are easily recognizable by the reddish-orange shoulder-spot and
the dark brownish-olive colour of the upper parts. From the
remarks of Berlepsch and Taczanowski it is evident that the
specimens from Huambo, N. Peru, belong to the same race.

(b) A. AURANTITROSTRIS ERYTHRORHYNCHUS Scl.

Arremon erythrorhynchus Sclater, P. Z.S. 1855, p. 83, pl. lxxxix.
(June 1855—~‘in Nova Grenada, Bogota”); idem, 1c. p. 154
(Bogota) ; idem, ].c. 1858, p. 83 (Bogota).

A. spectabilis Sclater, Cat. B. Brit. Mus. xi. p. 276 (part. a, b:
Bogota).

Habitat. Eastern Corompi4: known only from Bogota
collections.

Characters. Back, larger upper wing-coverts, and outer webs
of remiges clear yellowish olive-green; bend of the wing light
lemon-yellow ; bill small and stout (like the preceding form).
Wing: ¢ 74, 2 70; tail: ¢ 60-64, 2 56-58; bill 13 mm.

Material. A series from Bogota in the Munich Museum.

Observations. Bogota skins are quite uniform in their characters
and always readily distinguishable.

(c) A. AURANTIIROSTRIS OCCIDENTALIS, subsp. n.

A. erythrorhynchus (nec Sclater) Sclater, P.Z.S. 1860, p. 85
(Nanegal, W. Ecuador); idem, ].c. p. 274 (Babahoyo); idem,
]. c. p. 293 (Esmeraldas); Salvadori & Festa, Boll. Mus. Zool.
Torino, xiv. no. 357, 1899, p. 21 (Rio Peripa, Gualea, W. Ecuador ;
crit., synon.).

A. spectabilis (nec Sclater) Taczanowski, P.Z.8. 1877, p. 332
(Palmal, distr. of Guayaquil, S.W. Ecuador); Sclater & Salvin,
P.Z.S. 1879, p. 505 (Remedios, W. Colombia): Berlepsch &
Taczanowski, P.Z.S. 1883, p. 548 (Chimbo, S.W. Ecuador) ;
Sclater, Cat. B. Brit. Mus. xi. p. 275 (part.: ec, d, Remedios ; i—n,
Sta. Rita, Nanegal, Babahoyo, W. Ecuador); Hartert, Nov. Zool.
v. 1898, p. 483 (Cachabi, Chimbo); Goodfellow, Ibis, 1901,
p. 471 (Santo Domingo, 8. Nicolas); Ménégaux in: Mission
Are Merid. Equat. ix. 1, p. B 81 (S. Domingo, Pachijal).

Habitat. Western Ecuapor: Palmal (Jelski), Babahoyo
(Fraser), Chimbo (Stolzmann, Rosenberg), Santa Rita (Buckley),
Gualea, Rio Peripa (Festa), Santo Domingo (Goodfellow, Rivet),
Nanegal (Fraser), Esmeraldas, (Fraser), Pachijal (Rivet), Cachabi
(Rosenberg). etc. WESTERN CoLompra: Juntas, Condoto, Rio
San Juan, Chocé (Palmer), Remedios, Antioquia (Salmon).

Characters. Colour of back, larger wing-coverts, and outer webs
of quills intermediate between A. a. spectabilis and A. a. erythro-
rhynchus, neither dark brownish olive nor clear yellowish green,
but intense olive-green, somewhat shaded with golden: bend of
the wing lemon- or gamboge-yellow, but never orange; bill

1120 MR. C, E. HELLMAYR ON THE

considerably larger, longer as well as slenderer, than in both of its
allies—Wing : ‘S 702 75, 2 67-70; tail: g 58-63, 2 53-58;
bill 14-153 mm.

Type in the Zoological Museum, Munich, no. 09.5862. ¢ ad.
Condoto, Rio Condoto, W. Colombia, 150 feet, 27 March, 1909.
M. G. Palmer coll. no. 2573.

Material. Two adult males from Western Colombia, eight
adults (of both sexes) and young from Western Hcuador (Quito,
Paramba, San Javier, Lita, Carondelet, etc.).

Observations. I cannot perceive any difference between Co-
lombian and Ecuadorian specimens. The characters indicated
above are perfectly constant in the large series (15) examined
by me.

A. a. occidentalis ranges from near sea-level up to about

3500 feet (Paramba, Rio Mira).

54, PsrrTosPIzA RIEFFERII RIEFFERII Boiss.

Tanagra Riefferit Boissonneau, Rey. Zool. 11. p. 4 (1840.—
Santa-Heé-de- Bogota, Colombia).

Psittospiza riefferi Sclater & Salvin, P. Z.8. 1879, p. 505
(Envigado, Concordia, Medellin, Remedios, Santa Elena).

Nos. 2790, 2791. ¢ Q@ ad. Tatama Mountain, 7600 ft, 9.x.09.
Wing 110, 106; tail 81, 79; bill 18 mm.

‘“‘ tris brown, feet and bill scarlet.”

Agreeing with specimens from Bogota and Western Ecuador,
but bill shghtly larger.

P.r. riefferii is well-known as an inhabitant of the mountains
of Colombia and Keuador. In Peru and Bolivia it is replaced
by P. riefferii elegans Tsch.*, of which we have a good series
from various localities in the Munich Museum.

55. SALTATOR ATRIPENNIS Scl.

Saltator atripennis Sclater, Proc. Acad. N. Sei. Philad. vii.
p. 261 (1856.—Popayan, Colombia); Sclater & Salvin, P. Z.5.
1879, p. 505 (Medellin).

No. 2821. ¢ ad. Pueblo Rico, 5200 ft., 29.x.09.— Wing 108;
tail 102; bill 19 mm.

“Tris dark brown, feet and bill black.”

This bird, which can be considered as typical, agrees perfectly
with Sclater’s original description, having the top of the head black
etc. ‘Two specimens from the neighbourhood of Quito (Kcuador)
in our Museum have a somewhat shorter, stouter bill, and the
crown dark cinereous, merely mottled with blackish on the forehead
and above the white eyebrow. However, they may be not
quite adult.

S. atripennis is known only from the highlands of Colombia
and Heuador.

* Saltator elegans Tschudi, Arch. f. Nature. 10; i. D. 288 (1844.—Peru).

BIRDS OF WESTERN COLOMBIA. 21

56. Savraror MAximus P. L. S. Miill.

Tanagra maxima P. L. 8. Miller, Natursyst. Suppl. p. 159
(1776—ex Daubenton, Pl. Enl. 205: Cayenne).

Saltator magnus Sclater & Salvin, P. Z.S. 1879, p. 505 (Reme-
dios, Medellin, Neche); Berlepsch, Journ. f. Orn. 1884, p. 293
(Bucaramanga).

Nos. 2432, 2434. $9 9 ad. Novita: 24.x11.08—Wing 100,
98; tail 89,91; bill 18, 19 mm.

Nos. 2195, 2021. ¢ 2 ad. Noanama: 27.viii., 17.x.08.—Wing
OS 9a- tal) 92. So) bill) 19 mm:

Nos. 2383, 2047. 9 2 imm. Noéovita: 9.xi1.09; Noanama:
2.1x.08.

“Tris dark brown, feet dark grey, bill black, mandible blue-
grey.”

The majority of these skins have the back decidedly purer and
lighter green than our large series from Eastern South America
(Cayenne, Venezuela, Para, E. Ecuador, etc.), though one or two
are scarcely distinguishable on this score.

57. Piryitus erossus Linn.

Loxia grossa Linneus, Syst. Nat. xu. 1, p. 307 (1766.—
“ America”: ex Brisson; we substitute Cayenne as type locality).

Pitylus grossus Cassin, Proc. Acad, Philad. 1860, p. 140 (Falls
of the River Truando); Sclater & Salvin, P.Z.S. 1879, p. 505
(Remedios, Neche).

Nos. 2114, 2157, 2158. g gad. Sipi: 26.ix
93-97 ; tail 82-85; bill 18-19 mm.

No. Q115. 2. Sipi: 24.ix.08.—Wing 93; tail 83; bill 174 mm.

« Tris dark brown, feet grey, bill bright scarlet.”

Not different from specimens obtained in Venezuela, Brazil,
and Eastern Ecuador.

., 0.x.08.— Wing

58. CACICUS UROPYGIALIS Lafr.

Cassicus uropygiali s Lafresnaye, Rev. Zool. vi. p. 290 (1843.—
** Colombie,” sc. Santa Fe de Bogots) ; Sclater & Salvin, P. Z.S.
1879, p. 509 (Jerico).

C. uropygialis ¢ Cassin, Proc. Acad. N. Sci. Philad. 1860, p. 139
(Falls of the R. Truando).

Nos. 1995, 2009, 2088. ¢ g ad. Mouth of Hate Viviane
Noanama: 25.vill.; near Sipi: 17.1x.08.—Wing 133-138; oil
96-98 ; bill 28-30 mm.

Nos. 1996, 2092. 9 9 ad. Mouth of Calima: 17.viil.; Sipi:
19.ix.08.—Wing 122, 123; tail 90; bill 26, 28 mm.

“Tris light (turquoise) blue, feet black, bill light yellow.”

The series agrees with Bogots skins and others from N.W.
Ecuador. C. macrorhynchus Scl. & Salv.*, from Central America,

* Cassiculus microrhynchus Sclater & Salvin, P. Z.S. 1864, p. 353 (Lion
Hill, Panama).

WARY MR. C. E. HELLMAYR ON THE

is much smaller and has a considerably shorter, slenderer bill, but
is most probably only the northern representative of C. uro-
pygialis. The specimens obtained by Lieut. Michler’s party
on the Falls of the Rio Truando, although referred by
Mr. Ridgway * to C. microrhynchus, are more likely to belong
to the present form (¢/7. Cassin, Proc. Ac. N. Sci. Philad. 1867,
p. 64).

59, AMBLYCERCUS HOLOSERICEUS Licht.

Sturnus holosericeus Lichtenstein, Preis-Verz. Mex. Vog. p. |
(1831.—Mexico) ; Journ. f. Ornith. xi. 1863, p. 55 (veprint).

No. 1982. (2). Guineo, Rio Calima: 8.vii.08.—Wing 92;
tail 86; bill 273 mm.

‘Tris white, feet black, bill light yellow.”

This specimen agrees with others from Western Ecuador, in
size and colour. I have not seen typical Mexican birds, which
may be different.

60. CASsIDIX ORYZIVORA VIOLEA Bangs.

[Oriolus oryzivorus Gmelin, Syst. Nat. 1, 1. p. 386 (1788—ex
Latham : Cayenne) ].

Cassidix oryzivora violea Bangs, Proc. New Engl. Zool. Cl. u.
p- 11 (1900.—La Concepcion, Santa Marta, Colombia).

Cassidix oryzivora (errore) Sclater & Salvin, P. Z.S. 1879,
p. 510 (Concordia, Antioquia, Remedios) ; Sclater, P. ZS. 1860,
p- 140 (Pallatanga, W. Ecuador) ; Salvadori & Festa, Boll. Mus.
Zool. Torino, xiv. no. 357, p. 30 (Rio Peripa, R. Daule,
W. Ecuador); Hartert, Nov. Zool. v. p. 485 (Paramba, Chimbo,
W. Ecuador).

No. 2429. g ad. Noévita, 150 ft., 23.x11.08.—Wing 200;
tail 152; bill 37 mm.

No. 2823. g ad. Pueblo Rico, 5200 ft., 30.x.08—Wing 210;
tail 162; bill 393 mm.

No. 2413. 9 ad. Noévita: 18.x11.08. Wing 153; tail 120;
bill 34 mm.

“Tris light yellow, bill and feet black.”

C. o. violea is an excellent form, easily recognizable by the
strong violet-purple gloss of the plumage. In typical (. o.
oryzivora, from Cayenne, N. Brazil, Venezuela, etc., the general
colour is dull black with bronzy reflection, and very little, if any,
violet admixture on pileum and chest. Specimens from Western
Eeuador aiso belong to C. 0. violea.

61. XANTHORNUS MESOMELAS SALVINII Cass.

[ Psarocoliuse mesomelas Wagner, Isis, 1829, p. 755 (1829.—
Mexico.)|

Teierus Salvinii Cassin, Proc. Acad. N. Sci. Philad. 1867, p. 51

* Bull. U.S. Mas. no. 40, pt. ii. p. 190.

BIRDS OF WESTERN COLOMBIA. 1723

(1867.—Costa Rica, Nicaragua, New Granada (Atrato River,
Bogot), ete.; type from Costa Rica, ¢fr. Stone, tc. vol. li.
1899, p. 34).

I. msomelas Cassin, Proce. Ac. Philad. 1860, p. 140 (River
Atrato) ; Sclater & Salvin, P. ZS. 1879, p. 509 (Neche).

No. 2614. 2 ad. Condoto, R. Condoto: 20.iv.09.—Wing 94 ;
tail 100; bill 20 mm.

No. 1971. S$ juv. Guineo, R. Calima : 5.vii1.08.

“Tris dark brown, feet blue, bill black.”

The adult bird agrees with Bogota skins. The length of the
wing, in the latter, varies from 93 to 96 mm. None of my
Colombian specimens showing any trace of white edging to the
inner secondaries, I consider XY. m. salviniias a well-characterized
race.

In 8.W. Ecuador (district of Guayaquil) and N.W. Peru
(Tumbez, Lechugal, Callacate, etc.) another distinct form is met
with. It has been named _Y. m. taczanowskii Ridgw.*, and may
be recognized by its inferior size (wing 85-88 mm.), lesser extent
of black at the base of the outer rectrices, and by having con-
spicuous yellowish-white edges to the innermost secondaries.

Cfr. Berl. & Tacz. P. Z.S. 1883, p. 552 (s. n. 7. mesomelas).

62. HypopyRRHUS PYROHYPOGASTER Tarragon.

Cassicus pyrohypogaster Tarragon, Rev. Zool. x. p. 252 (1847.—-
“ Nouvelle Grenade ”).

Hypopyrrhus pyrrhogaster Sclater & Salvin, P.Z.S. 1879,
p- 510 (Envigado, Santa Elena).

Nos. 2832, 2833. 9 2 (in moult). Pueblo Rico, 5200 ft.,
4.xi.09.

No. 3768. ¢ ad. Siaté, 5200 ft., 23.ix.09.—Wing 142;
tail 140; bill 31 mm.

Nos. 3758, 3761, 3762, 3770. 29 9. Siaté: 17,18, 23.1x.09.—
Wing 127-130; tail 128-133; bill 27-29 mm.

“Tris light yellow, feet and bill black.”

The specimens agree with a number of Bogota skins in the
Munich Museum. As will be seen from the above measurements
the females have much shorter wings and tail.

H. pyrohypogaster is vestricted to the highlands of Colombia. It
is not uncommonly met with in Bogota collections, and Salmon
procured several examples in Antioquia. |

63. CYANOCORAX AFFINIS AFFINIS Pelz.

Cyanocorax affinis Pelzeln, Sitzungsber. Ak. Wiss. Wien,
math.-naturw. Cl. xx. 1, p. 164 (1856.— Bogota); Sclater &
Salvin, P. Z. 8. 1879, p. 510 (Cauca, Remedios).

* Teterus mesomelas taczanowskii Ridgway, Proc. Wash. Acad. Sci. iii. p. 153
(1901.—Guayaquil, 8. W. Ecuador).

WA Bat MR. C. E. HELUMAYR ON THE

C. pileatus (nec Temminck) Cassin, Proc. Acad. N. Se. Philad.
1860, p. 138 (Rs. Truando and Nercua, N.W. Colombia *).

Nos. 2053, 2459. g¢ 2 ad. Noanama: 3.1x.08, 11.1.09.—
Wing 170, 165 ; ; tail 175, 168; bill 27, 29 mm.

No. 2537, So ‘afl El Tigre, R. Tamana, 13.11.09.— Wing 165 ;
tail 166; bill 29 mm.

No. 3746. g imm. Pueblo Rico, 5200 ft., 10.1x.09.—Wing
170; tail 160; bill 29 mm.

‘Tris lemon-yellow, feet and bill black.”

In the pale yellowish-white colour of the under parts and tail-
end these birds agree with Bogota skins, but are slightly smaller 7
In size they approach C. a. zeledoni, from Chiriqui and Costa me
the latter form may, however , easily be distinguished by the bright
creamy-yellow belly, ete.

A good series from Carthagena (the type locality of C. sclateri
Heine {) is required to show whether the shght difference in size
Hee birds from Bogota and Western Colombia is constant.

C. a. affinis is confined to Colombia, ranging from near sea-
level a | to about 5000 feet altitude.

64. CYANOoLYcA PULCHRA Lawr.

Cyanocitta pulchra Lawrence, Ann. Lye. N. H. N.Y. xi. 1875,
p. 163 (Feb. 1876.—Quito, Ecuador).

No. 2789. gad. Tatama Mountain, 7600 ft., 9.x.08.— Wing
130; tail (Gmeulting) about 135; bill 830 mm.

“Tris, feet, and bill black.”

So far as I am aware this is the second known specimen of
this fine species, which was described by Mr. Lawrence from a
single Quito skin in his collection. The bird corresponds exactly
to the original deseription with the sole exception that the smoky-
blackish chest shows a faint cobalt-bluish tinge. C. pulchra cannot
be confused with any other species. It differs from all the forms
of the C. armillata group by the’ beautiful silvery bluish-
white colour of the crown and nape, terminated behind by a
distinct cross-band of bright ultramarine blue; by the dull
brownish-black upper back, without any blue tinge; by the much
duller cobalt-blue throat-patch; by lacking the black jugular
crescent, so conspicuous a feature in all of its allies; by the smoky-
blackish chest, and much duller bluish belly, etc. Moreover, the
bill is much stouter and larger, while the tail, on the other hand,
is much shorter.

There can be no doubt that C. pulchra is an excellent species,
not much like any other member of the genus.

* Ridgway (Bull. U.S. Mus. no. 50, pt. ili. p. 305), with a query, refers the above
quotation to the Central American race C. a. zeledoni Ridgw. (Auk, xvi. 1899,
p. 255.—Talamanea, Costa Rica). But as specimens from Carthazena, N. Colombia.
agree with C. a. affinis from Bogota ete., those from Truando and Nercua most
certainly belong also to the latter, inasmuch as the fauna of those districts is
essentially the same as that of the San Juan River.

+ Bogoté skins measure: wing 175-180; tail 175-180 mm.

¢ Journ. f. Ormith. vii. p. 114 (1860. — Cartagena (an in Guatemala),”.

BIRDS OF WESTERN COLOMBIA. 1125

65. SAYORNIS CINERACEA CINERACEA Lafr.

Tyrannula cineracea Lafresnaye, Rev. Zool. xi. p. 7 (1848.—
“ Caracas, in Venezuela”).

Sayornis cineracea Sclater & Salvin, P.Z.S8. 1879, p. 511
(Medellin, Frontino).

S. ardosiacus (nec Tyrannula ardosiaca Lafresnaye, 1844)
Cassin, Proc. Acad. N. Sc. Philad. 1860, p. 144 (Falls of the Rio
Truando); efr. Ridgway, Bull. U. 8. Mus. no. 50, pt. iv. pp. 512,
(footnote c), 594 (note a).

No. 2831. @ imm. Pueblo Rico, 5200 ft., 4.xi1.09.— Wing 81 ;
tail 73; bill 145 mm.

“Tris dark brown, feet and bill black.”

Tdentical with examples from Merida and Bogota.

66. Copurus LEUCONOTUS Lafr.

Copurus leuconotus Lafresnaye, Rev. Zool. vy. p. 835 (1842—

“in Bolivia,” errore ; we substitute Bogota, Colombia *); Sclater
& Salvin, P. Z.S. 1879, p. 511 (Remedios).

Nos. 1983, 2054, 2068, 2280, 2346. 5 gad. Gnineo; 10.vi1i.08 ;
Noanama: 3, 5.ix.08 ; Novita: 13, 28.xi.08.—Wing 72-75; tail
170-183 ; bill 8-9 mm.

Nos. 1984, 2345, 2347. 9 9. Guineo: 10.viii.08.; Novita;
27, 28.xi.08.— Wing 70; tail 125-135; bill 8mm.

“ Tris dark brown, feet and bill black.”

The females are smaller, with the median rectrices less elongated,
and the pileum is uniform dark sooty without hoary margins.

There is apparently no difference between specimens from so
widely separated localities as Costa Rica and Colombia, nor do
skins from Western Heuador differ.

67. PLATYTRICCUS MYSTACEUS ALBOGULARIS Scl.

[Platyrhynchos mystaceus Vieillot, Nouv. Dict. xxvul. p. 14
(1818—ex Azara: Paraguay). |

Platyrhynchus albogularis Sclater, P. Z.S. 1860, p. 68 (1860.—
Pallatanga, W. Ecuador).

P. cancroma (nec Temminek) Cassin, Proc. Acad. N. Sci. Philad.
1860, p. 144 (Truando).

No. 2811. g ad. Pueblo Rico: 25.x.09.—Wing 62; tail 36 ;
bill 12 mm.

“Tris dark brown, feet pink, bill black.”

This bird is quite typical. Skins from Chiriqui and Costa
Rica agree in coloration, but are slightly smaller (wing 5/—
59 mm.). I have not seen specimens from Eastern Ecuador 7 and
Northern Peru { which may, or may not, be strictly referable

* “ Bolivia” is an obvious pen-slip for “Colombia,” the paper having as title
“ Oiseaux nouveaux de Colombie.”

+ Tacz. & Berl. P. Z.S. 1885, p. 88 (Machay).

* Taczanowski, P. Z.S. 1879, p. 233 (Tambillo).

Proc. Zoot. Soc.—1911, No. LX XVI. 76

1126 MR. C E. HELLMAYR ON THE

to P. m. albogularis. Birds from Northern Venezuela (San
Esteban, Tocuyo, etc.), however, are very different, being much
paler and more greenish brown above, having the throat tinged
with pale yellowish, the lower mandible whitish instead of
mainly blackish, etc., ete. ‘They agree perfectly with topotypes of
P. insularis Allen* from Tobago, and other examples from
Trinidad and Bermudez (Cumana), while a series secured by the
late H. Whitely on Mount Roraima, Guiana, is intermediate
between P. m. mystaceus (of 8.E. Brazil and Paraguay) and
P.m. insularis. Cfr. Hellmayr, Nov. Zool. xiii. 1906, p. 22.

According to my views, the range of the various races of the
group is as follows :—

(a) P. mystaceus mystaceus Vieill. South-eastern Brazil
from Bahia and Minas southwards to Rio Grande do Sul,
Paraguay, and the adjoining province Misiones, Argentine.

(b) P. mystaceus insularis Allen. Tobago, Trinidad, and North
coast of Venezuela t, from Cumand (Bermudez) west to
Tocuyo. [Specimens from Mt. Roraima, British Guiana,
and from the Orinoco R. (Caicara), may form yet another
race. |

(c) P. mystaceus albogularis Scl. Western Keuador and
Colombia, Panama, Chiriqui, and parts of Costa Rica.
[East Ecuador and North Peru 2]

68. RHYNCHOCYCLUS CINEREICEPS FLAVoTECTUS Hart.

[Cyclorhynchus cinereiceps Sclater, Ibis, i. p. 443 (1859.—
Oaxaca, Mexico). |

Rhynchocyclus megacephala flavotectus Hartert, Nov. Zool. 1x.
p. 608 (1902.—San Javier, N.W. Ecuador).

Nos. 2250, 2396. 3 g ad. Novita: 8.xi., 12.xii.08.— Wing 63 ;
tail 503, 51; bill 12, 13 mm.

No. 2046. dg imm. Noanama: 2.ix.08.—Wing 62; tail 51;
bill 12 mm.

“Tris brown, feet dark grey, maxilla black, mandible horn-
coloured.”

The specimens agree In every respect with the types from San
Javier and two other examples from Paramba, N.W. Ecuador,
kindly lent by the authorities of the Tring Museum. ‘The species
is for the first time recorded from Colombia. 2. ¢. flavotectus,
although described as a subspecies of 2. megacephala auct. nec
Swains.{, is much more nearly allied to 2. cinereiceps, of Central
America, in fact its southern representative. In the coloration
of the under parts the two birds are perfectly alike; but
R. c. flavotectus may be recognized by its much duller and darkec

* Platyrhynchus insularis Allen, Bull. Amer. Mus. N. Y. ii. p. 143 (1889.—
Tobago).

+ P. albogularis Sclater, Cat. B. xiv. p. 67 (part., specimens d, e: S. Esteban,
Venezuela).

{ Cfr. Hellmayr, Verhandl. zool.-bot. Ges. Wien, liii. 1903, pp. 206-8.

BIRDS OF WESTERN COLOMBIA, WN D7/

green (instead of yellowish-green) back, much brighter yellow edges
to the remiges, and by having the median and greater wing-coverts
broadly edged with bright yellow. As a rule, the cinereous
colour of the pileum is somewhat darker, and the tail on the
average shorter.

Specimens from N.W. Ecuador measure as follows :—

(S)ad. §S. Javier: wing 66; tail 53; bill 12mm.

(2)ad. 8. Javier (type): wing 62; tail 51; bill 12 mm.

Two adults (“¢, 2”), Paramba: wing 62, 63: tail 50, 513;
bill 124, 13 mm.
_ Lat first thought that 2. sulphurescens asemus Bangs * might be
the same as the present species. However, Mr. Bangs gives much
larger dimensions (wing 70, tail 59 mm.) and compares his new
form with R&. sulphurescens, to which R. c. flavotectus is but very
distantly related.

R.c. flavotectus is another characteristic form’of the forest district
of the Pacific Coast, ranging from sea-level up to about 3500
feet (Paramba).

69. CRASPEDOPRION ZQUINOCTIALIS Scl.

Cyclorhynchus ceguinoctials Sclater, P. Z. 8. 1858, p. 70
(1858.—Rio Napo, HE. Keuador).

C. brevirestris (nec Cabanis) Cassin, Proc. Acad, N. Sci. Philad.
1860, p. 144 (Rio Truando).

Rhynchocyclus equinoctialis Hartert, Nov. Zool. v. 1898, p. 487
(Cachabi, N.W. Ecuador).

No. 2422. gad. Novita: 21.xii.08—Wing 78; tail 65; bill
16 mm.

No. 2485. gd ad. Noanama: 16.109.—Wing 80; tail 69;
bill 15 mm.

“Tris dark brown, feet dark blue, maxilla black, mandible
horny-white.”

These skins are practically identical with an adult male from
N.W. Ecuador. All have the greater upper wing-coverts and the
inner secondaries conspicuously edged with ochreous-buff, exactly
like C. olivaceus Temm.,f from Eastern Brazil; they differ,
however, from the last named-species in their larger size, much
larger bill, deeper green throat and chest, etc.

Tam not sure whether they are really referable to C. equinoctialis,
not having examined specimens from Eastern Ecuador (the type
locality). According to Sclater’s original description and later
remarks, the true C. wquinoctialis appears to lack the fulvous
margins to the wing-coverts and inner secondaries.

C. cequinoctialis, as understood here, ranges from Ecuador
north to the Isthmus of Panama.

* Proc. Biol. Soc. Wash. xxiii. p. 73 (1910.—Pavas, Western Colombia, 4400 ft.).

+ Platyrhynchos olivaceus Temminek, Rec. Pl. Col., livr. 11. pl. 12. fig. 1
(1820.—* Brésil ”).

t Cat. B. Brit. Mus. xiv. p. 166.
76*

1128 MR. C. E. HELLMAYR ON THE

70. TopIROSTRUM CINEREUM CINEREUM Linn.

Todus cinereus Linneus, Syst.. Nat. x11. 1, p. 178 (1766—ex
Edwards, Glean. N. H. ii. p. 110, pl. 262. fig. inf.: Surinam).

Todirostrum cinereum Cassin, Proc. Acad. N. Sci. Philad. 1860,
p. 144 (Carthagena); Sclater & Salvin, P.Z.S. 1879, p. 512
(Medellin, Santa Elena, Remedios).

No. 1973. ¢ ad. Guineo, Rio Calima: 6.vii.08— Wing 44;
tail 35; bill 14 mm.

‘“‘ Tris white, feet blue-grey, bill black.”

Perfectly identical with specimens from Bogota, Venezuela
(San Esteban), Cayenne, etc. Chin and throat are yellow lke the’
remaining under parts.

In Western Ecuador this form is represented by Jodirostrum
cinereum sclateri Cab. & Heine*, at once known by having the
chin and upper throat white. It extends southwards into the
district of Tumbez, N.W. Peru.

71. CNIPODECTES SUBBRUNNEUS SUBBRUNNEUS Scl.

Cyclorhynchus subbrunneus Sclater, P. Z. 8. 1860, p. 282
(1860.—Babahoyo, 8.W. Ecuador).

Cnipodectes subbrunneus Sclater & Salvin, P. Z. 8. 1879,
p- 514 (Remedios, Antioquia).

No. 2509. SG ad. Névita: 30.1.09.—Wing 94; tail 87; bill
17 mm. .

No. 2539. g ad. Hl Tigre: 13.11.09—Wing 95; tail 92;
bill 17 mm.

Nos. 2605, 2606. ¢ g ad. Condoto: 16, 16.iv.09.—Wing 95,
92; tail 89, 86; bill 164, 17 mm.

No. 2591. S ad. Condoto: 10.iv.09.—Wing 87; tail 83;
bill 163 mm. :

No. 2569. 3 imm. Névita: 15.1.09.—Wing 75; tail 69; bill
163 mm.

“Tris pink (in adults), brown (in imm. male, no. 2569),
feet grey, maxilla black, mandible pink (adults), horn-coloured
(young).”

This series is most interesting, proving as it does that the so-
called “ minor” from Panama and Western Kcuador are nothing
but the young of the larger swbbrunnews. This had already
been suspected by me when, some years ago, I carefully studied and
measured the entire material in the British Museum, but in the
absence of a good series from any locality, it was then impossible
to arrive at a definite conclusion. The occurrence side by side of
these two ‘‘ species” differing only in size always appeared to
me a phenomenon hardly reconcilable with modern views of
zoogeographical distribution.

No. 2569, a young bird, as manifested by the fluffy texture of
the plumage, differs from the other specimens in its much shorter

* Triccus Sclateri Cabanis & Heine, Mus. Hein. ii. p. 50 (1859.—Peru).

BIRDS OF WESTERN COLOMBIA. 1129

wings and tail, less developed crest, more rufescent brown colour
of the upper parts, and especially in having distinct, ochraceous-
buff apical edges to the greater wing-coverts. An unsexed bird
obtained at Santa Rita, W. Ecuador, by Buckley’s collector
Villagomez, and two skins, one marked “ ¢,” from Panama,
“ McLeannan leg.,” in the British Museum, are also immature.
They were erroneously referred by Dr. Sclater to the Peruvian
race C'. s. minor, the type of which is likewise a young bird !
However, adult males from Chamicuros (the type locality of
C. minor) differ slightly from those of Western Ecuador and
Colombia (Remedios) in having both upper and under surfaces
vather paler brown. An adult male obtained on the Rio Puris,
W. Brazil, shows the same variation when compared with our
Chocd series. Hence the name C. s. minor may well be
retained for the Amazonian race, though its (rather trifling)
characters should be confirmed by a better series of fresh skins.
According to my views there are two very closely allied races :—

(1) 0. s. subbrunneus Scl., ranging from 8.W. Ecuador
through Western Colombia to Panama.

(2) OC. s. minor Scl., ranging from Eastern Peru (Chami-
curos) to W. Brazil (Rio Purts).

Their synonymy and ranges are as follows :—

(a) C. SUBBRUNNEUS SUBBRUNNEUS Scl.

Cyclorhynchus subbrunneus Sclater, P. Z. 8. 1860, p. 282
(1860.—Babahoyo, 8.W. Ecuador) ; idem, 1. c. p. 295 (Hsmeraldas,
N.W. Ecuador); Lawrence, Ann. Lyc. N. H. N.Y. vii. 1862,
p. 473 (Panama).

Myiochanes subbrumneus Sclater, Cat. Amer. B. 1862, p. 232
(Babahoyo, Esmeraldas).

Myiochanes sp., Sclater & Salvin, P. Z.S. 1864, p. 360 (Panama).

Cnipodectes subbrunneus Sclater & Salvin, P. Z.S8. 1879, p. 514
(Remedios, Antioquia); Sclater, Cat. Birds Brit. Mus. xiv.
p. 197, pl. xvi. (part.: Babahoyo, Esmeraldas, Balzar: Ecuador ;
Remedios, Colombia); Salvin & Godman, Biol. Centr.-Amer., Aves,
ii. 1889, p. 55 (part.: Colombia, Panama, Ecuador); Salvadori &
Festa, Boll. Mus. Torino, xiv. no. 362, 1899, p. 10 (Rio Peripa,
W. Ecuador); Ridgway, Bull. U. 8. Mus. no. 50, pt. iv. 1907,
p- 484 (Panama, Colombia, W. Ecuador).

C. minor (nec Sclater) Sclater, Cat. B. Brit. Mus. xiv. p. 197
(part. : d, Santa Rita, W. Ecuador; e,/ Panama); Ridgway, Bull.
U.S. Mus. no. 50, pt. iv. p. 485 (part.: Panama, W. Ecuador).

Type locality. Babahoyo, 8.W. Ecuador.

Range. Western Ecuador: Babahoyo, Esmeraldas (Fraser), Rio
Peripa (Festa), Balzar (Jilingworth), Santa Rita (Villagomez).
Western Colombia: Novita, El Tigre, Condoto, Choco distr.
(Palmer); Remedios, Antioquia (Salmon). Panama: Lion Hill
(MacLeannan), Cascajal Coclée (7eyde).

1130 MR. GC. BE. WELLMAYR ON THE

Material.
Wing. Tail. Bill.
Brit. Mus.—One adult from Babahoyo (type)............. 86; 82; 17° mm.
» Aj One adult from Esmeraldas, Ecuador ... 92; 82; Wee op
55 - One adult from Balzar, Ecuador ............ 853 86; LGin ites

Munich Mus.—Five ¢ g ad. from Choeo, W.Colombia. 87-95; 83-92 ; 165-17
Brit. Mus.—One ¢ ad. from Remedios, W. Colombia. 88; 83 ; 17
3s a One juv. from Sta. Rita, Ecuador ......... 74; 72; 16
Pr Two juv. from Panama ......................... 71,735 65,68; 15, 16
Manic Mus.—One ¢ juy. trom Chocd, ive (Caltrain. 75; 69; Goes
Adults from W. Ecuador and Colombia are practically
identical. From Panama I have seen only young birds.

(6) C. SUBBRUNNEUS MINOR Scl.

Cnipodectes ninor Sclater, P. Z.S. 1883, p. 654 (1884.—
Chamicuros, E. Peru); Taczanowski, Orn. Pérou, ii. 1884,
p. 294 (Chamicuros); Sclater, Cat. B. Brit. Mus. xiv. p. 197
(part.: a—c, Chamicuros); Ridgway, Bull. U.S. Mus. no. 50,
pt. iv. 1907, p. 485 (part.: Eastern Peru).

Cnipodectes subbrunneus minor Suethlage, Journ. f. Orn. 1908,
p- 12 (Rio Purts, Brazil),

C. subbrunneus (nec Sclater) Sclater & Salvin, P. Z.S8. 1873,
p. 281 (Chamicuros); Salvin & Godman, Biol. Centr.-Amer., Aves,
1. 1889, p. 55 (part.: Peru); Berlepsch & Leverkuhn, Ornis vi,
1890, p. 16 (Chamicuros).

Type locality. Chamicuros, Eastern Peru.

Range. Kastern Peru: Chamicuros (Bartlett), W. Brazil:
upper R. Purus (Zuber).

Material.
: Wing. Tail. Bill.
Mus. Brit.—Two adult males from Chamicuros ......... 92; 84,83; 153mm.
» s One ¢ juv. from Chamisuros (type) ...... 72; 64; NWS | sy
Mus. Paré.—One adult male from Rio Purvis ............ 86; q7 Gies

72, SERPOPHAGA CINEREA CANA Bangs.

[Huscarthmus cinereus Styickland, Ann. Mag. Nat. Hist. xiii.
p. 414 (1844.—* Chili”’).]

Serpophaga cinerea cana Bangs, Proce. Biol. Soc. Wash. xvii.
p- 113 (1904.—Chirua, Sierra de Santa Marta, Colombia).

S. cinerea (nec Strickland) Sclater & Salvin, P. Z. 8. 1879,
p. 512 (Envigado, Frontino).

No. 2741. g ad. Pueblo Rico, 5200 ft., Sept. 1909.— Wing 564 ;
tail 47; bill 9 mm.

“Tris dark brown, feet and bill black.”

Identical with specimens from Bogotéand Mérida (Venezuela).
Birds from Southern Peru (Andes of Carabaya) which I take
to be true cinerea are darker grey above, and have the breast
more strongly shaded with cinereous, while those from Costa
Rica and Chiriqui, S. cinerea grisea Lawr.*, are smaller, with the
crown more deeply black.

* Serpophaga grisea Lawrence, Ann. Lyc. N. H. N.Y. x. p. 189 (1871.—San
José, Costa Rica).

BIRDS OF WESTERN COLOMBIA. 1131

73. MIONECTES OLIVACEUS HEDERACEUS Bangs.

[ Mionectes olivaceus Lawrence, Ann. Lye. N. H. N. Y. ix.
p- 111 (1868.—Barranea and Dota, Costa Rica). ]

Mionectes oleagineus hederaceus Bangs, Proc. Biol. Soc. Wash.
xxii. p. 73 (1910.— Pavas, Western Cordillera, Colombia,
4400 ft.).

M. striaticollis (evrore) Selater, P. Z. 8. 1859, p. 144 (Palla-
tanga, W. Ecuador); idem, P. Z. 8. 1860, p. 93 (Nanegal, W.
Ecuador); Berlepsch & ‘Taczanowski, P. Z. S. 1884, p. 296
(Surupata, Chaguarpata, W. Ecuador; jide H. v. Berlepsch
in litt.).

M. olivaceus (errore) Hartert, Nov. Zool. v. 1898, p. 487
(Chimbo); Salvadori & Festa, Boll. Mus. Zool. Torino, xiv.
no. 362, 1899, p. 6 (Niebli, Rio Peripa, W. Ecuador) ; Goodfellow,
Ibis, 1901, p. 704 (San Nicolas, Gualea, Canzacota, W. Ecuador) ;
Hartert, Nov. Zool. ix. 1902, p. 607 (Lita, Paramba, N.W.
Keuador).

Nos. 1960, 1998. og. San Joaquim, Bahia del Chocé,
l.viii.08; Boca de Colima, R. San Juan, 18.viii.08.—Wing 68,
69; tail 54; bill 123, 13 mm.

Nos. 922565 2263, 2322) 6 dads Novita: LO 21.08-—
Wing 67-69; tail 58; bill 12-13 mm.

No. 2252. Q ad. Novita: 9.xi.08—Wing 62; tail 48; bill
12 mm.

“Tris brown, feet light grey, maxilla black, base of mandible
pale brown.”

Palmer’s series as well as several examples from Western
Keuador differ at a glance from true JZ, o. olivaceus, of Costa Rica
and Chiriqui, in having the back much duller, less yellowish
green, the pileum more greyish green, and the olive streaks on
the anterior under parts much darker and more strongly marked.
The Colombian and Venezuelan races, WV. o. galbinus Bangs* and
M. o. venezuelensis Ridgw.t, are much brighter green above, the
pileum being scarcely darker than the back, and the general colour
of the lower surface is much deeper yellow, with the dark streaks
more decidedly green.

M. 0. hederaceus replaces the typical form in the lowlands and
on the Pacific slopes of the Western Cordillera of Colombia and
Ecuador.

Whether its ally Jf. striaticollis D’Orb. & Lafr. also occurs
in Western Colombia ¢ I am unable to ascertain, but it should be
mentioned that the numerous skins I have examined from
Western Ecuador were all referable to J/. 0. hederaceus.

* Mionectes olivaceus galbinus Bangs, Proc. New Eng. Zool. Cl. iii. p. 85
(1902.—La Concepcion, Santa Marta, N. Colombia).

+ M. olivaceus venezuelensis Ridgway, Proc. Biol. Soc. Wash. xix. p. 116
(1906.—Guacharo, State of Cumana, N. Venezuela).

{ M. striaticollis Sclater & Salvin, P. Z.S. 1879, p. 512 (Santa Elena).
: IL. s. poliocephalus Bangs, Proc. Biol. Soc. Wash. xxiii. 1910, p. 74 (San Antonio,

io Cali).

WS MR. C. E. HELLMAYR ON THE

74, LEPTOPOGON SUPERCILIARIS POLIOCEPHALUS Cab. & Heine.

| Leptopogon superciliaris Tschudi, Arch. f. Naturg. 10, i. p. 275
(1844.—Peru, 7. e. “ Vorwiilder” of Central Peru; ¢/7. Tschudi,
Faun. Peru., Aves, p. 162.|

L. poliocephalus Cabanis & Heine, Mus, Hein. ii. p. 55 (1859.—
** New Granada,” sc. Bogota).

L. superciliaris (nee Tschudi) Hartert, Nov. Zool. ix. 1902,
p. 607 (Lita, Paramba, N.W. Heuador).

Yo. 2376. dad. Nodovita, 150 ft., 7.x11.08.— Wing 69 ; tail 62;
bill 15 mm.
“Tris dark brown, feet and bill black.”

A second specimen from this region I have seen in Count
Berlepsch’s collection. It is an adult female, and was obtained
by the late Gustav Hopke at San Pablo, 4500 ft. alt., January 21,
1897. (Wing 65; tail 583; bill 143 mm.)

The West Colombian birds agree with a series from Bogota
and San Esteban, Venezuela, in size and coloration. Specimens
from N.W. Ecuador (Paramba) are also similar.

All these examples differ obviously, however, from ZL. s. swper-
celiaris, of which I have examined a large series from Central and
Southern Peru and Northern Bolivia, in the much deeper yellow
abdomen, and in having the throat and chest strongly tinged with
olive-green. The typical race has the under parts very much
paler, about primrose-yellow, with but a slight greenish wash on
breast and sides.

The colour of the apical spots to the wing-coverts is of no diag-
nostic value, for it varies from primrose-yellow to dull ochreous
in individuals taken at the same place,

While there canbe no question as to the distinctness of Z. s
superciliaris and L. s. poliocephalus, the status of the third
geographical race, ZL. s. transandinus Berl. & Tacz.*, from
South-western Ecuador, is not yet definitely settled. Two females
in Count Berlepsch’s collection are, indeed, much smaller than
LL. s. poliocephalus ; but the dimensions given J. c. for a male are
scarcely inferior to the average measurements of adult males
from Colombia and Venezuela, . So far as coloration is concerned
the two forms are practically identical.

Another uncertainty exists with regard to the Central Ameri-
ean birds, which Ridgway and other American naturalists have
referred to ZL. s. superciliaries of Peru. This, however, can
hardly be correct. Hither they belong to JZ. s. poliocephalus,
ov they may represent yet another race.

75. KLaNIA CINEREA PARAMB& Hellm.

| Elainea cinerea Pelzeln, Zur Orn. Bras. ii. pp. 108, 180 (Sept.
1860.— Marabitanas). |

* P. Z. 8. 1883, p. 553 (1884.—Chimho, S.W. Ecuador).

BIRDS OF WESTERN COLOMBIA. 4133

Serpophaga parambe Hellmayr. Bull. B. O. C. xiv. p. 54 (1904.—
Paramba, N.W. Ecuador).

[Tring Museum. ¢ ad. Paramba, N.W. Ecuador, 3500 ft.
23.vii. 1899. Miketta coll. no. 473, Type of species.— Wing 565 ;
tail 48; bill 10 mm.|

No. 2494. d imm. Névita: 26.1.08.—Wing 534; tail 44 ;
bill 95 mm.

No, 2196. ¢ juv. Noanama: 17.x.08.—Wing 56; tail 49;
bill 10 mm.

“ Tris dark brown, feet and bill black, base of mandible brown.”

On receipt of these specimens it at once occurred to me that
they might represent some phase of S. paramba, originally
described from an adult male in the Tring Museum; and the
careful examination of the typical example not only confirmed
the correctness of my surmise, but, furthermore, clearly showed
that S.parambe is merely a western, smaller subspecies of Hlania
cinerea.

In fact, on comparing the types of the two “species” I find
them identical in structural characters as well as in coloration.
However, /. c. parambe may be recognized by its much shorter
wings and tail, and by its rather slenderer, shorter bill.

The immature males obtained by Mr. Palmer differ from the
type in the following points:—The back is bright olive-green
instead of bluish grey; the tips to the upper wing-coverts and
edges to the secondaries are pale yellow, instead of white, the
edges to the rectrices olive-green, not cinereous. Moreover,
the throat only is white, while the remaining under parts,
including axillaries and under wing-coverts, are pale yellow, with
obsolete, greyish-white flammulations on the chest. |The vertical
patch, in no. 2494, is white with a hardly perceptible yellowish
hue in its posterior portion, the rest of the pileum ashy grey, as in
the adult male, but slightly darker.

The immature birds are not unlike the corresponding stage of
F. c. cinerea*, but can easily be distinguished by having the
pileum ashy-grey (not olive-green like the back), the coronal patch
white (instead of pale yellow), and the under parts much paler
yellow, with the throat conspicuously white. The bill is also
narrower and shorter.

FE. c. parambe: evidently replaces H.c. cinerea on the western
sides of the Andes in Ecuador and Colombia.

For comparison, the dimensions of seven skins of /. c. cinerea.
are herewith given :—

Five adult males +... . Wing 60-63 ; tail 55-57; bill 11-12 mm.
Two immature birds ¢ OE O0 40-03 5 0) ye te mtn

* Described by me in Nov. Zool. xv. 1908, p. 46.

+ (a) Marabitanas, Rio Negro: type of species; (4) Bogoté, Mus. H. v. B.;
(c) Sarayacu, Eastern Ecuador, type of S. albogrisea Scl. & Salv.; (d) Chamicuros,
FE. Bartlett coll.; (¢) Suapure, Caura, Tring Museum.

t (a) ? jr. Suapure, Caura, Tring Museum; (4) Bogota, Mus. Berlepsch.

1134 MR. C. E. HELLMAYR ON THE

76. Luaarus AL3ICOLLIS ALBICOLLIS Vieill.
Tyrannus albicollis Vieillot, Nouv. Dict. xxxv. p. 89 (1819—

ex Azava: Paraguay).

Nos. 2648, 2658. ¢ g ad. Tad6, 230 ft.: 17, 21.v.03—Wing
82, 81; tail 58, 59; bill 11, 12mm.

Nos. 2654 (not numbered). 2 2 juv. Tado: 18, 19.v.09.—
Wing 78; tail 58, 59; bill 11 mm.

“Tris brown, feet and bill black.”

Not different from Brazilian specimens.

77. MYIoZErETES CAYANENSIS CAYANENSIS Linn.

Muscicapa cayanensis Linneeus, Syst. Nat. xii. 1, p. 327 (1766—
ev Brisson: Cayenne).

Hlenia cayennensis Cassin, Proc. Acad. N. Sci. Philad. 1860,
p. 144 (Turbo).

Myiozetetes texensis (errore) Sclater & Salvin, P.Z.S. 1879,
p-. 513 (Hnvigado, Medellin).

No. 2384. Q ad. Novita: 9.xi11.08.—Wing 83; tail (@ncom-
plete); bill 135 mm.

No. 1992. ¢ jr. Bocade Calima: 4.viii.08.—-Wing 85; tail 73 ;
bill 14 mm.

“Tris dark brown, feet and bill black.”

The adult bird agrees with other specimens from Bogota,
Cayenne, and Pard. About the geographical races of this species
cfr. Hellmayr, Abhandl. Bay. Akad. Wiss. Miinchen, i. Cl. xxi. 3,
1906, pp. 649-650.

M. c. rufipennis Lawr. *, synonymised with the typical form by
Mr. Ridgway‘, is easily recognizable by the greater extent of
rufous on the wings. It inhabits the northern littoral of Vene-
zuela (Valencia, Puerto Cabello, ete.) and is not to be confounded
with MM. c¢. erythroptera Latr.¢ from South-eastern Brazil
(Rio, Southern Minas Geracs), which is much larger and has
still more rufous on the wings.

78. CoryPHOrRICCcUS ALBOyITTATUS Lawr.

Pitangus albovittatus Lawrence, Ibis, iv. p. 11 (1862.—Isthmus
of Panama).

Coryphotriccus albovittatus Ridgway, Bull. U.S. Mus. no. 50,
pt. iv. 1907, p. 669 (Panama).

No. 2421. g ad. Novita, Rio Tamana: 21.x11.08.—Wing 81 ;
tail 69; bill 17 mm.

“Tris brown, feet and bill black.”

C. albovittatus, one of the rarest of Neotropical Tyrants, is
apparently recorded for the first time from South America, it

* Myiozetetes rufipennis Lawrence, Ann. Lyc. N. H. N. Y. ix. p. 267 (1869.—
Valencia, Venezuela).

y7 Bull. U.S. Mus. no 50, pt. iv. p. 446.!

£ Lyrannula erythroptera Latresnaye, Rey. Mag. Zool. (2) y. p. 56 (1853.—Brazil).

BIRDS OF WESTERN COLOMBIA. 1135

having previously been known only as an inhabitant of Teeter
Panama.*

The bird agrees very welll with Ridgway’s description except
for its slightly larger size 7 and the colour of the pileum, which
I should call sooty- -blackish rather than sooty-brown. From
C. parvus Pelz. = it differs in its duller, less greenish back and
white throat G@nstead of yellow like the remaining under parts).
In structural characters the two species are per fectly alike.

So far as I know, the above is the only specimen existing in any
European Museum.

79. MyioB1lus SULPHUREIPYGIUS VILLOSUS Scl.

[Tyrannula sulphureipygia Sclater, P. Z.S. 1856, p. 296 (Jan.
1857.—Cordova, Mexico). |

Myiobius villosus Sclater, P. Z. S. 1860, P 93 (1860.—
Nanegal, W. Ecuador) ; Selater & Salvin, P. 7.8 . 1879, p. 514
(Fr nen).

M. sulphureipygius (errore) Cassin, Proc. Acad. N. Sci. Philad.
1860, p. 144 (Rio Truando).

No. 2427. g ad. Névita: 22.xii.08.—Wing 63; tail 54; bill
11 mm.

‘‘Tris dark brown, feet grey, maxilla black, mandible pink.”

This bird, like others from Western Ecuador, differs from
M. s. aureatus Bangs §, of Chiriqui and Costa Rica (Pozo Azul),
in its darker, more brownish-green back, deeper ochraceous chest
and sides, and more brownish under tail-coverts. The differences
between the two races, though slight, appear to be quite constant.
Mr. Ridgway || makes J/. sulphureipygius a subspecies of MZ. aan-
thopygus | =mastacalis Wied |, but this is certainly wrong.

M. b. mastacalis is merely the southern form of J/. barbatus,
the two being completely connected by intermediates in North-
eastern and Northern Brazil (Piauhy,- Maranhao, Para, Rio
Madeira). The representative of the barbatus group in Western
Kceuador, Colombia, Panama, and Costa Rica is J. 6. atricaudus
Lawr., which is clearly specifically distinct from JZ. s. villosus,
with which it occurs in many localities. Mr. Rosenberg’s cor-

respondents sent numbers of both forms from the same places
in N.W. Ecuador.

* Whether the newly described C. a. distinctus Ridgw. (Proc. Biol. Soc. Wash.
xxi. 1908, p. 191—Rio Reventazén, near Guaydbo, E. Costa Rica) is really distinct
can, of course, only be decided when series trom Panama and Eastern Costa Rica
become available.

+ Ridgway gives for Panama specimens the following measurements : wing, ¢
79s, 2 Wi tails Gas, 2 663 bill) 16; 162, mm:

f£ Pitangus. parvus Pelzeln, Zur Ornith. Bras. ii. pp. 111, 181 (1868.—Marabi-
tanas, Rio Negro).

§ WM. xanthopygus aureatus Bangs, Proc. New Engl. Zool. Cl. iv. p. 27 (1908.—
Chiriqui).

Il Bull. U.S. Mus. no. 50, pt. iv. p. 490. *

* Cfr. Hellmayr, Abhandl. Meat Wiss. Miinchen, ii. Cl., xxii. 3, 1906, pp. 641-3.

1136 MR. C. E. HELLMAYR ON THE

Therefore, the natural affinities of these species are correctly
expressed by the following scheme :—

(a) WM. barbatus barbatus: Guiana and Northern Brazil (Rio
Negro, Borba, Para, etc.).

(b) I. barbatus atricaudus; Western Ecuador and Colombia,
Panama, Costa Rica.

(c) WM. barbatus mastacalis: Kastern Brazil from Bahia to Rio,
west to the Rio Madeira (cfr. Nov. Zool. xvii. 1910,
p- 299).

(d) MW. sulphureipygius sulphureipygius : Southern Mexico ;
2 Guatemala, British Honduras.

(ce) M. sulphureipygius aureatus: From Honduras south to
Panama.

(f) M. sulphureipygius villosus : Western Colombia and Ecua-
dor; ? Peru, N. Bolivia.

80. MyIoBIUs ERYTHRURUS FULYIGULARIS Saly. & Godm.

[Myiobius erythrurus Cabanis, Arch. f. Naturg. 13) ap 249°
pl. 5. fig. 1 (1847.—Guiana, Cayenne). |

M. fulvigularis Salvin & Godman, Biol. Centr.-Amer., Aves,
ii. p. 58 (1889.—Santa Fe, Veragua).

No. 2554. 9 ad. Juntas, Rio Tamana, 23.11.09.—Wing 48 ;
tail 36; bill 84 mm.

“Tris brown, feet yellow, maxilla black, mandible pink.”

This is an extreme example of I. e. fulvigularis, agreeing with
skins from Costa Rica. ‘The forehead is strongly washed with
buff, the under parts are bright buffy ochreous, the throat hghter,
more tawny, etc.

Ofr. my remarks in Nov. Zool. xiv. 1907, p. 48.

81. EMPIDONAX VIRESCENS Vieill.

Platyrhynchos virescens Vieillot, Nouv. Dict. xxvii. p. 22
(1818—based on Muscicapa querula (nee Vieillot) Wilson, Amer.
Orn. ii. 1810, p. 77, pl. 13. fig. 2—Eastern North America).

Empidonax acadicus auct. (nec Gmelin).

No. 2826. (¢)ad. Pueblo Rico: 5200 ft., 2.xi.09.—Wing 74 ;
tail 62; bill 13 mim.

No. 2174. @ ad. Sipi: 9.x.08.—Wing 70; tail 59; bill
12 mm.

“ Tris brown, feet dark grey, maxilla black, mandible yellow.”

A common winter visitor to the Western States of South
America. It has several times been taken in Hcuador.

82. MyI0CHANES RICHARDSONII RICHARDSONII Swains.

Tyrannula richardsonit Swainson, Fauna Bor.-Amer. 11. p. 146,
pl. 46, fig.inf. (1831.—Cumberland House, Saskatchewan, Canada).

No. —. @ ad. Loma Hermosa, 4150 ft.: 22.x.09.— Wing 80;
tail ‘62 ; bill 12 mm.

BIRDS OF WESTERN COLOMBIA. WN S7

“Tris dark brown, feet and bill black.”

This species is also a regular winter visitor to Western South
America, The specimen sent by Mr. Palmer agrees with others
from Costa Rica (San José), Quito, etc.

83. MyzArcuus crinitus Linn.

Lurdus crinitus Linneus, Syst. Nat. x. p. 170 (1758—ex
Catesby, Carolina, i. p. 52, pl. 52: Carolina).

Myiarchus crivitus Berlepsch, Journ. f. Ornith. 1884, p. 303
(Bucaramanga).

Nos. 2409, 2420. 6 ¢ ad. Novita: 17, 21.x11.08.— Wing 101,
107 ; tail 92, 96; bill 21, 22 mm.

No. 2394. 9 (2) ad. Novita: 12.xi1.08.— Wing 1062 ; tail 98 ;
bill 21 mm.

No. 2469. g¢ ad. Noanama: 13.1.09.—Wing 110; tail 102;
bill 22 mm.

“Tris dark brown, feet black, maxilla black, mandible horn-
brown.”

These birds are absolutely identical with skins from the Eastern
United States. From MW. tyrannulus (P. L. 8. Mull.) they differ
in being decidedly greenish instead of greyish brown above, in
the darker cinereous throat and chest, much deeper yellow belly,
and in having the whole of the inner web of the outer rectrices
rufous.

The present record extends the range of MM. crinilus in its
winter quarters considerably to the south. The most southerly
locality yet known was Bucaramanga, whence Lorentz had sent
a specimen to the Bremen Museum (cfr. Berlepsch, 1. c. p. 303).

84. MyYIARCHUS NIGRICEPS Scl.

Myiarchus nigriceps Sclater, P. Z.S. 1860, p. 68 (1860.
tanga, W. Ecuador).

No. 2809.(¢)ad. Pueblo Rico, 5200 ft.: 25.x.09.—Wing 82 ;
tail 75; bill 18 mm.

No. 2208. ¢ imm. Noanama: 21.x.08.—Wing 77; tail 73;
bill 18 mm.

“Tris dark brown, feet and bill black.”

The two skins agree with topotypical examples from Western
Keuador. The pileum is uniform deep black.

Mr. Ridgway, in his great work *, has again confounded two
distinct species under the name J/. nigriceps, overlooking my
remarks in Nov. Zool. xiii. 1906, p. 23 & pp. 323-4, I think I
have conclusively shown that the birds from Venezuela, Amazonia,
and Guiana belong to J. tuberculifer, while M. nigriceps is
restricted to Panama, W. Colombia, W. Ecuador, and Northern
Peru.

The range of the two species is given 7m eatenso in Noy. Zool.
xiii. p. 26.

Palla-

* Bull. U.S. Mus, no. 50, pt. iv. p. 650.

1138 MR. C. E. HELLMAYR ON THE

85. TYRANNUS MELANCHOLICUS SATRAPA Cab. & Heine.

[Tyrannus melancholicus Vieillot, Nouv. Dict. xxxv. p. 84
(1819.—ex Azara: Paraguay). |

Laphyctes satrapa Cabanis & Heine, Mus. Hein. ii. p. 77
(1859.—* Guiana and Caracas”).

Tyrannus melancholicus (nec Vieillot) Sclater & Salvin, P. Z.S.
1879, p. 516 (Retiro, Medellin) ; Cassin, Proc. Acad. N. Sci.
Philad. 1860, p. 143 (Turbo, Carthagena, R. Truando).

No. 2712. g ad. Tadé6, 230 ft., 17.vi.09.—Wing 116; tail
101; bill 21 mm.

“ Tris dark brown, feet and bill black.”

85. CHLOROPIPO HOLOCHLORA Lira Hellm.

[Chloropipo holochlora Sclater, Cat. B. Brit. Mus. xiv. p. 287
(1888.— Bogoti). |

Chloropipo holochlora lite Hellmayr, Nov. Zool. xiii. p. 325
(1906.—Lita, N.W. Ecuador).

CO. holochlora (nec Sclater) Sclater, Cat. B. Brit. Mus. xiv.
p- 287 (part.: specimen 6, “ Pasto,” Colombia); Hartert, Nov.
Zool. v. 1898, p. 488 (Cachabi, Paramba, N.W. Ecuador).

No. 2081. g ad. Sipi: 12.x.08.—Wing 69; tail 44; bill
103 mm.

No. 2352. (3) ad. Novita: 30.%1.08:—Wing 71; tail 46;
bill 113 mm.

No. 2112. 9 ad. Sipi: 23.1x.08.—Wing 663; tail 43; bill
11 mm.

“Iris brown, feet grey or blackish, maxilla black, mandible
grey.”

Identical with topotypical skins from North-western Eeuador.
The upper parts are light olivaceous green, the throat, chest, and
sides somewhat duller. On the eastern (Amazonian) slope of
the Colombian and Ecuadorian Andes this form is replaced by
typical C. h. holochlora.

The locality ‘ Pasto” requires confirmation. The specimen is
typical of C. h. lite.

O. h. lite is restricted to the lowlands and foot-hills of
W. Colombia and N.W. Ecuador.

87. MAsIUS CHRYSOPTERUS BELLUS Hart. & Hellm.

[Pipra chrysoptera Lafresnaye, Rev. Zool. vi. p. 97 (1843.—
Bogota). |

Masius chrysopterus bellus Hartert & MHellmayr, Orn.
Monatsber. xi. p. 35 (1903.—Riolima, Cauca Valley, 4000 ft.).

M. coronulatus (nec Sclater) Sclater & Salvin, P.Z.S. 1879,
pp- 516, 549 (Antioquia ; no exact locality); Pelzeln & Madarasz,
Monogr. Pipr. pl. v. (figure of the Antioquia specimen).

No.—. ¢ ad. Pueblo Rico, 5200 ft., 28.x.08.— Wing 59; tail
43; bill 7 mm.

BIRDS OF WESTERN COLOMBIA. 1139

“Tiis brown, feet light red, maxilla light grey, mandible pale
red.”

This specimen confirms the characters of the subspecies origin-
ally based upon a male obtained by the late J. H. Batty in the
same region. ‘The feathers of the coronal crest, which have
exactly the same shape as in Jf. ¢. coronulatus of Western
Eeuador, being thickened at the end into a horny substance,
are dark brownish red instead of light tobacco-brown.

M.c. bellus is well figured in Pelzeln and Madarasz’s Monograph
of the Pipride, on plate v. In their joint communication *
Hartert and Hellmayr have pointed out that Salmon’s Antioquia
specimen belongs to this form, calling attention to a misleading
statement in Sclater and Salvin’s report in P. Z.8. 1879. AV. .
bellus is as yet known only from the Western Cordillera of
Colombia.

88. PIPRA MENTALIS MINOR Hart.

[Pipra mentalis Sclater, P. Z.S. 1856, p. 299, pl. 121 (1857.
— Cordova, Vera Cruz, Mexico). |

Pipra mentalis minor Hartert, Nov. Zool. v. p. 489 (1898.—
Cachabi, N.W. Ecuador). |

No, 2225. g ad. Rio Cajén: 31.x.08.—Wing 60; tail 29;
bill 8 mm.

No. 2098. d ad. Sipi: 21.1x.08.—Wing 60; tail 29; bill
85 mm.

Nos. 2002, 2201. ¢ g ad. Noanama: 22.viii, 19.x.08.—Wing
58, 57; tail 28, 29; bill 8, 8 mm.

Nos; 220%, 2308, 23805 2371. 3g Sg ad. Novita: 185 Oia;
5, 9.xi1.08.—Wing 57-58 ; tail 28-29; bill 8 mm.

Nos. 2030, 2192, 2430. g S juv. (plumage of female).
Noanama: 27.vill.; Sipi: 14.x.; Novita: 23.x11.08—Wing 60;
tail 29-303 ; bill 84-9 mm.

Nos. 2172, 2226.2 Qad. Sipi: 9.x.08; RioCajon: 31.x.08.—
Wing 59; tail 28, 303; bill 83-9 mm.

“Tris white (in adult male), dark brown (in females and young
males), feet light brown or grey, maxilla dark brown, mandible
pale brown or pink.”

The adult males agree, so far as coloration is concerned, with
typical birds from Northern Ecuador. The head is of the same
intense crimson hue, the gonydeal angle is but narrowly edged
with yellowish, and the imner secondaries only have slight
yellowish-white margins along the inner web. While the bill is
as small and short as in Ecuador specimens, in the length of the
wing and tail the Colombian birds approach P. m. ignifera,
from Chiriqui and Costa Rica. The difference in size, however,
is very slight and not constant enough to warrant their separation
from P. m. minor. Ten adult males from Western Ecuador

* Ornithologische Monatsberichte, xi. pp. 33-25.

1140 MR. C. E. HELLMAYR ON THE

(including type) measure as follows :—wing 55 (twice) to 58; tail
26-28 ; bill 8-9 mm.

P.m. minor thus ranges from Ecuador northwards over the
Pacific lowlands of Western Colombia.

89. PrpRA CORONATA VELUTINA Berl.

| Pipra coronata Spix, Av. Bras. 11. p. 5, pl. vii. fig. 1 (1825.—
S. Paulo d’Olivenca). |

Pipra velutina Berlepsch, Ibis, 1883, p. 492 (1883.—Veragua).

P. cyaneocapilla (nec Hahn) Sclater & Salvin, P.Z.S, 1879,
p- 517 (Medellin, Remedios, R. Neche, Antioquia).

Nos. 2301, 2316, 2340, 2365, 2401. ¢ g ad. Névita: 17, 20,
26.x1., 3, 15.xn-08.

Nos. 2118, 2105, 2146, 2156, 2182, 2241. g g ad. Sipi: 22,
Mae sscegy Ila he 8). 14 oe ONS)

No. 2030. ¢ ad. Noanama: 26.x1.08.

No. 2239. g ad. Rio Cajon: 4.xi.08.

Nos. 2031, 2040, 2058, 2216. g g juv. Noanama: 28, 31.viii.,
Ae OO.

No. 1964. g juv. 8S. Joaquim: 4.viii.08. No. 2160. 3 juv.
Sipi: 6.x.08.

Nos. 2311, 2334, 2361, 2362, 2366. 9 9 ad. Néovita: 19
24.x1., 1, 3.x11.08.

Nos. 2121), 2124, 2173. @ 2 ad. (Sipi: 25, 28.1x, 9.x.08:

No. 1965. 9 ad. 8S. Joaquim: 4.v11.08.

“Tris dark red (in adult males), dark brown (in females and
young males), feet black, maxilla black, mandible blue.”

3d ad.: wing 554-58; tail 24-26; bill 7-8 mm.

@ 2 ad.: wing. 55-58; tail 25-28; bill 74-85 mm.

In ‘The Ibis’ for 1906, pp. 31-32, I have already alluded to
the smaller size of specimens from N.W. Ecuador and Western
Colombia. All of the many skins procured by Mr. Palmer have
much smaller bills and decidedly shorter wings than a series from
Chiriqui and Costa Rica. In the adult males the black of the
plumage is also somewhat deeper. Although I feel pretty sure
that the southern birds constitute a separable race, yet I should
like to examine a series from Veragua, the type locality, before
making any formal separation.

Specimens from different localities measure as follows :—

?

Males. Wing. Tail. Bill.
mm. mm mm.
Oe nt GUN I acs cee oe aa staiee 62-64 27-29 9-92
ALSO NW 5. CORE) ARO incoodounsgdooAes 62-64 27-28 9-93
3: Panama (Railway) ....:...... 58-59 26-27 1-74
N4bo WY, Gollommlon® soacascscceacsoscc 509-08 243-26 7-8
TS INN 5 WGC GoaShoaeanaddne 5d-57 =. 25-27 = 74-8
Females.
4: Chiriqui and Costa Rica...... 614-63 28-30 93-10
LO Wis Colom nia i eeerereeeneeeetee 544-57 25-27 74-8
SING IBCURNTOIE: Luscusscendsae HG BYE 7-8

BIRDS OF WESTERN COLOMBIA. 114)

90. CHIROMACHERIS VITELLINA Gould.
Pipra vitellina Gould, P. Z. 8. 1843, p. 103 (Dec. 1843,—

Panama).

Chiromacheris vitellina Sclater & Salvin, P..Z.8. 1879, p. 516
(Cauca, Remedios).

Nos. 2025, 2097, 2364, 2494. ¢ g¢ ad. Noanama: 28.viii;
Sipi: 2l.ix; Novita: 3.xii.08, 27.1.09.— Wing 53-54; tail 28-30 ;
bill 10 mm.

Nos. 2067, 2185. 3 gjuv. Noanama: 5.1x.08; Sipi: 13.x.08.

Nos. 2352, 2439. 9 9 ad. Noéovita: 30.x1., 28.x1.08.— Wing
54-55 = taal 29; bill 10 mm.

“Tris dark brown, feet orange, bill black ”

A single male from Panama is rather more yellowish green on
the belly, but does not differ otherwise. C. awrantiaca Salv. is
apparently the geographical representative of C. vitellina in
Veragua and 8.W. Costa Rica.

C. vitellina ranges from the Isthmus of Panama south to
W. Colombia.

91. SapAYoA =NIGMA Hart.

Sapayoa enigma Hartert, Nov. Zool. x. p. 117 (1903.—Rio
Sapayo, N.W. Ecuador); Hellmayr in Wytsman’s Genera Avium,
part ix. 1910, p. 28, pl. 1. figs. 5, 8.

No. 2418. gad. Noévita: 19.xi1.08—Wing 82; tail 66;
bill 15 mm.

No. 2070. g juv. Noanama: 1.1x.08.—Wing 81; tail 61;
bill 135 mm.

Nos. 2071, 2457. 9 imm., 9 ad. Noanama: 5.ix.08, 9.1.09.—
Wing 82, 79; tail 59; bill 15 mm.

“Tris brown, feet black, bill black, mandible grey in adult male
and females, yellow in the young male” (no. 2070).

This curious bird was hitherto represented only by a single
female, the type, from N.W. Keuador, in the Tring Museum.
Tt has been fully described by Hartert, and I have nothing to add
to his excellent account. Sapayoa is certainly related to
Scotothorus, though the shape of the bill, the feathering of the
upper portion of the metatarsus ete., serve to distinguish it at a
glance. ;

The females agree exactly with the type, while the adult male
differs in having the middle of the crown bright golden yellow.
The tail is longer, with the rectrices decidedly narrower, and
the central pair slightly elongated, exceeding the submedian one
by about 3mm. The immature female has the upper part of
the head more yellowish green, and the throat slightly paler
yellowish than the adults. The young male (without any trace of
the golden yellow crown-patch) has a much shorter, smaller bill,
with the lower mandible yellowish white instead of brownish
horn-colour. The range of S. enigma is restricted to the humid
lowlands of the Pacific coast of N.W. Ecuador and W. Colombia.

Proc. Zoo, Soc.—1911, No. LX XVII. ca

1142 MR, C. E. HELLMAYR ON THE

92. 'TITYRA SEMIFASCIATA COLUMBIANA Ridgw.

| Pachyrhynchus semifasciatus Spix, Av. Bras. 11. p. 32, pl. xliv.
fig. 2 (1825.—Para). |

Tityra semifasciata columbiana Ridgway, Proc. Biol. Soc.
Wash. xix. p. 119 (1906.—La Concepcion, Santa Marta).

T. personata (nec Jardine & Selby) Sclater & Salvin, P. Z. 8.
1879, p. 517 (Remedios, Neche).

No. 2502. g ad. Noévita: 29.1.09—Wing 122; tail 77;
bill 24 mm.

No. 2314. 9 ad. Névita: 19.x1.08—Wing 114; tail 69;
bill 24 mm.

“Tris red-brown (¢ ), dark-brown (Q ), feet dark grey, bill red,
tip black.”

As correctly pointed out by Mr. Ridgway, the adult male differs
from typical 7’. semifasciata of Amazonia * by the much
narrower black frontal area, the posterior margin of which ends
in a line with the middle of the eye, instead of extending as far as
its posterior edge. Moreover, the black band across the inner
web of the outer rectrices is much broader, and reaches nearly or
quite to the shaft. The female is much browner on the upper
parts, with the pileum considerably darker.

By the coloration of the female, and the greater extent of
black on the lateral rectrices, this form approaches 7’. s. costaricensis
Ridew. + from Panama and Costa Rica, which, however, is
smaller and has even more black in the tail.

T. s. columbiana ranges from Santa Marta through Western
Colombia to the province of Hsmeraldas, N.W. Ecuador. Speci-
mens from the latter locality agree in every respect with the
Colombian ones.

93, TrryRA ALBITORQUES ALBITORQUES Dubus.

Vityra albitorques Dubus, Bull. Acad. Roy. Belg. xiv. pt. 11.
p. 104 (1847.—Peru); Sclater & Salvin, P. Z. 8. 1879, p. 517
(Remedios).

Nos. 2278, 2452. gg ad. Novita: 13.xi1.08; Noanama:
7.1.08.—Wing 107, 105; tail 67; bill 214, 225 mm.

No. 2525. S ad. El Tigre, R. Tamana: 10.11.09.— Wing 103 ;
tail 64; bill 22 mm.

No. 2524. 9 ad. El Tigre: 9.11.09.—Wing 100; tail 64;
bill 22 mm.

‘“‘ Tris dark brown, feet grey, bill black.”

The males do not appear to be different from those obtained
by Natterer at Manaos, N. Brazil, or from an adult male from

* Mr. Ridgway (Bull. U. S. Mus. no. 50, pt. iv. p. 868) gives as its range
“*Southern Brazil and Bolivia to Central Colombia.” ‘This is not quite correct,
for T. s. semifasciata is strictly an Amazonian species, ranging from Paré west to
the Eastern slopes of the Andes, south to H. Bolivia and W. Mattogrosso.

+ Proc. Biol. Soe, Wash. xix. p, 119 (1906,—Bonilla, Costa ica).

BIRDS OF WESTERN COLOMBIA. iat;

Chyavetas, N. Peru. The back, upper wing- and tail-coverts are
pale pearl-grey, the basal half of the rectrices greyish white, the
remaining portion black with a distinct white apical margin, the
ear-coverts white, etc., exactly as in the skins from Manaos and
Peru.

T. buckleyi Salv. & Godm.* from Eastern Ecuador, the type
of which I have examined in the British Museum, may be
distinguished by the paler, nearly white colour of the upper wing-
coverts and rump, by having the white in the tail confined
to the extreme base of the outer web of the external rectrices,
etc. ete.

94, PLATYPSARIS HOMOCHROUS Sel.

Pachyrhamphus homochrous Sclater, P. Z. S. 1859, p. 142
(1859.—Pallatanga, W. Ecuador).

Hadrostomus homochrous Sclater & Salvin, P. Z. 8. 1879, p. 517
(Remedios).

Nos. 2260, 2600, 2685. ¢ ¢ ad. Névita: 10.xi.; Condoto :
14.iv.09; Tadd: 2.vi.09.—Wing 90; tail 63-65; bill 16—-
17 mm.

No. 2415. $ juv. Novita: 19.x1.08.— Wing 86; tail 61;
bill 17 mm.

No. 2399. 9 ad. Novita: 17.xi1.08.—Wing 85; tail 61;
bill 17 mm.

“Tris dark brown, feet blue-grey, bill black, mandible grey in
the young male and female.”

Identical with skins from N.W. Ecuador, province of
Esmeraldas.

This species ranges from Panama (Railroad) and Santa Marta
in the north through Western Colombia and Western Hcuador
to N.W. Peru (Lechugal, district of Tumbez) 7.

95, PACHYRHAMPHUS DORSALIS Sel,

Pachyrhamphus dorsalis Selater, Cat. Amer. B. p. 245 (1862.—
““ Bogota ?” t); Ridgway, Bull. U. 8. Mus. no. 50, pt. iv. 1907,
p. 832 (Rio Cali, R. Barratoro, Castilla, W. Colombia); Bangs,
Proc. Biol. Soc. Wash. xxi. 1908, p. 157 (“ N.W. Colombia,
just south of Darien,” errore; see footnote above p. 1085),

P. cinerewentris (nec Sclater) Sclater & Salvin, P. Z. 8. 1879,
p. 518 (Santa Elena).

No. 3750. @. Pueblo Rico, 5200 ft., 11.ix.09.—Wing 75;
tail 57; bill 133 mm.,

‘“‘ Tris brown, feet dark grey, bill black.”

* Biol.-Centr, Amer., Aves, ii. p. 120 (Dec. 1890.—Yanayacu, Ecuador).

+ The locality “Sarayacu,” Hast Ecuador, is undoubtedly incorrect. I have, on
several occasions, alluded to the fact that Buckley’s localities cannot be relied upon,
collections from the western and eastern slopes of the Andes having been mixed
up and almost invariably labelled “Sarayacu.”

{ I have examined the type in the British Museum. [It is an undoubted Bogoté

skin, i
77*

1144 MR. C. E. HELLMAYR ON THE

I quite agree with Mr. Ridgway in considering this species
totally different from P. niger cinereiventris Scl., with which
Dr. Sclater later united it. In fact, P. dorsalis appears to be
the Western representative of P. marginatus Licht.* It may,
however, readily be distinguished in the male sex by the uniform
black upper back, separated from the glossy black pileum by a
very broad, light cinereous nape-band. The wings and tail are
longer, and the bill decidedly larger.

The female, as yet undescribed, is very much like that of
P. marginatus, but has a larger, broader bill, and the pilewm is
dark olive, each feather with an apical spot of dull greenish
black.

In addition to the type and another specimen in the British
Museum, I have examined four more adult males from Bogotd in
the collections of the Munich Museum and of Count Berlepsch.
The Tring Museum possesses specimens from Paramba, N.W.
Ecuador, which are perfectly similar to Bogota skins.

P. dorsalis thus ranges from Panama through Western
Colombia to N.W. Ecuador.

96. PACHYRHAMPHUS CINNAMOMEUS Lawr.

Pachyramphus cinnamomeus Lawrence, Ann. Lyc. N. H. N.Y,
vil. p. 295 (Jan. 1861.—Lion Hill Station, Panama. Railway).

Pachyrhamphus cinnamomeus Sclater & Salvin, P. Z. S$. 1879,
p. 518 (Remedios).

P. rufescens® Cassin, Proc. Acad. N. Sci. Philad. 1860, p. 189
(Turbo). ;

Nos. 1980, 2535. ¢ gd ad. Guineo: 8.vi1.08; El Tigre, 320 ft. :
12.11.09.—Wing 78, 79; tail 57, 55; bill 144 mm.

Nos. 1968, 2034. ¢ ¢ imm. (without the rudimentary second
primary). 8. Joaquim, Bahia del Choco, 4.viii; Noanama:
29.viii.08.— Wing 75, 79: tail 57; bill 14 mm.

Nos. 2026, 2350. 992 ad. Noanama: 28.vii.; Ndévita:
28.xi.08.— Wing 75, 76; tail 55, 58; bill 14 mm.

“Tris dark brown, feet blue-grey, maxilla black, mandible
blue.”

The series is exactly like specimens from Western Ecuador,
while skins from Central America (Costa Rica, Guatemala), as
a rule, are of a deeper rufous-tawny on the upper parts.

P. cinnamomeus ranges from Southern Mexico (Tabasco) and
Guatemala southwards to Western Hecuador ‘7.

97. LATHRIA UNIRUFA CASTANEOTINCTA Hart.

[|Lipaugus unirufus Selater, P. Z. 8. 1859, p. 385 (1860.—
Oaxaca, S. Mexico, and Guatemala). |

* =P. atricapillus auct. (nec Merrem); cf. Berlepsch, Nov. Zool. xv. 1908,
p. 141.

+ In Upper Amazonia (Eastern Ecuador; Ucayali, Peru) its place is taken by
another form, closely allied to, or perhaps even identical with, P. castaneus Jard.
& Selby. [= P. rufus auct, nec Boddaert !]

BIRDS OF WESTERN COLOMBIA. 1145

Lathria unirufus castaneotinctus Hartert, Nov. Zool. ix. p. 610
(1902.—Rio Durango, N.W. Ecuador).

Lathria unirufa (nec Sclater) Hartert, Nov. Zool. v. 1898,
p. +90 (Cachabi, N.W. Ecuador); Sclater & Salvin, P. Z.8. 1879,
p-. 918 (Remedios, Neche).

Lipangus unirufus (nec Sclater) Cassin, Proc. Acad. N. Sci.
Philad. 1860, p. 143 (Turbo, R. Truando).

Nos. 2306, 2315, 2045, 2175. ¢ gd ad. Novita: 18, 20.xi. ;
Noanama: 1.ix.08; Sipi, Rio Sipi: 10.x.08.—Wing 130-136 ;
tail 102-108; bill 20-21 mm.

Nos. 2917, 2342, 2343, 2423. 2 2 ad. | Cajon, Ry Cajon ;
28.x.08 ; Novita: All Sep 29, xii.08.— Wing 123-128; tail 101-105;
bill 20-21 mm.

“Tris dark brown, feet grey, maxilla brown, mandible pale
brown or grey.” !

These specimens agree with topotypical birds from N.W.
Ecuador and differ from Z. w. clara Ridgw.* of southern
Central America, in having the back decidedly brighter, more
cinnamon-rufous, and the under parts also darker, deep tawny-
ochraceous. The bill also is light enon instead of blackish
horn-colour.

The specimens from Antioquia (aomedion Neche) in the
British Museum likewise belong to this form; and so probably
also do those obtained by Lieut. Michler’s expedition to the Rio
Truando (although they are referred by Mr. Ridgway t to
Lu. clara), since the fauna of the Atrato region is essentially
identical with that of the Choc6 district.

LL. u. castaneotincta, therefore, appears to range from the coast
of the gulf of Uraba through Western Colombia to the province
of Esineraldas, N.W. Ecuador.

98. LIPAUGUS HOLERYTHRUS ROSENBERGI Hart.

[Lipaugus holerythrus Sclater & Salvin, P. Z. 8. 1860, p. 300
(1860.—Vera Paz, Guatemala). |

Lipaugus holerythr us rosenbergi Hartert, Bull. B. O. C. xvi. p. 12
(1905.—Rio Dagua, W. Colombia).

Lipaugus holerythrus (errore) Hartert, Nov. Zool. v. 1898, p. 489
(Cachabi, N.W. Ecuador).

Nos. 2355, 2375. 3 d ad. Néovita: 30.x1., 7 xii,08.—Wing 104,
NOD sian! 90, 895 bill 19, 20 mm.

No. 2100. 9 ad. Sipi, Rio Sipi: 21.ix.08.—Wing 102;
tail 90; bill 19 mm. ;

“Tris dark brown, feet dark grey or black, maxilla blackish,
mandible brown.”

These birds, which are to be regarded as topotypical of Z wu.
rosenbergi, differ from Central American skins in their darker

* Proc. Biol. Soc. Wash. xix. p. 120 (1906.—Lion Hill, Panama).
y+ Bull, U.S. Mus. no. 50, pt. iv. p. 824.

1146 MR. C. E. HELLMAYR ON THE

tawny russet upper, and deeper tawny under parts, as Sonal
pointed out by Hartert.

Although I have not seen the examples from Antioquia (Neche,
Remedios *), I have little doubt that they also are referable to
the present form rather than to typical L. h. holerythrus.

L. h. rosenbergi is another of the many species peculiar to
Western Colombia and N.W. Ecuador (province Esmeraldas).

99. CorinGA NATTERERID Boiss.

Ampelis nattererii Boissonneau, Rey. Zool. iii. p. 2 (1840.—
Santa-Fé-de- Bogota).

Cotinga nattererii Ridgway, Bull. U.S. Mus. no. 50, pt. iv.
1907, p. 785 (Hastern Panama (Railway line) to Colombia ;
crit.).

Cotinga simont Berlepsch, Ornis, xiv. p. 361 (1907.—San Jose,
Rio Dagua, W. Colombia).

Nos. 2267, 2268, 2269, 2281, 2283, 2289, 2290, 2291, 2292.
6 dad. Névita, Rio Tamang: 12 i 16.xi.08..-Wing 107—
111; tail 66-69; bill 143-153 mm.

No. 2270. gS imm. Novita: 12 ix.08.—Wing 108; tail 67;
bill 153 mm.

Nos. 2282, 2293. 6 g juv. Névita: 14, 16.xi.08.—Wing 105;
tail 68, 70; bill 14, 15 mm.

Nos. 2271, 2275, 2284, 2294, 2295, 2296. 9 9 ad. Novita:
12, 18, 14, 16.xi.08.—Wing 106-110; tail 71-75; bill 143-
15 mm.

“Tris black, feet dark grey, maxilla black, mandible blue.”

The series of adult males is fairly uniform, the variation
being chiefly confined to the intensity of the purple colour
of the throat and abdominal patch. Two specimens, especially
nos. 2281, 2291, have the throat darker than the others. All,
however, show the characteristic blackish cross-bars (whieh
sometimes have a slight metallic greenish sheen) to the feathers
of the throat, already noticed by M. Boissonneau and more fully
described by Mr. Ridgway, and the pale cerulean blue border to
the gonydeal angle; the pileum is constantly deeper blue than
the back; the second primary is very nearly as long as the third,
and never attenuated on its apical portion, so striking a feature
in the allied C. ridgwayi Ridgw., from Chiriqui and 8.W. Costa
Rica.

I am sorry to say that C. simoni is clearly a synonym of
C. nattererii, for the description of Boissonneau, which Count
Berlepsch appears to have overlooked, corresponds exactly to the
Chocé birds.

I do not understand the Count’s statement as regards the
coloration of the female which, he says, eens that of

* TL, holerythrus Sclater & Salvin, P. Z. S. 1879, p. 519 (Neche); Sc a er, Cat. B.
xiv. 1888, p. 357 (part.: Remedios, Neche).

BIRDS OF WESTERN COLOMBIA. 1147

C. cayana. As a matter of fact, the six females obtained by
Mr. Palmer are exactly like C. ridgwayi except in being slightly
deeper buff on the anterior under parts. In the main features,
viz., crown and back sooty edged with buffy white, larger
upper wing coverts broadly margined with cinnamon-buff, dusky
markings of lower surface etc., the females of the two species are
practically identical.

C. nattererii ranges from Eastern Panama (Railway line) south
to Buenaventura, Western Colombia. Occasionally it is also met
with in Bogota collections.

100. CARPODECTES HOPKE! Berl.

Carpodectes hopkei Berlepsch, Orn. Monatsber. v. p. 174 (1897.—
San José, R. Dagua, W. Colombia); Hartert, Nov. Zool. ix, 1902,
p-. 611 (Ventana, N.W. Ecuador).

Nos. 2205, 2272, 2285, 2286, 2287, 2288. ¢ ¢ ad. & imm.
Novita: 12, 14, 18.x1. 08.— Wing 158-170; tail 95-102; bill 20-
213 mm.

No. 2273. (¢) juv. Novita: 12.xi.08.—Wing 158; tail 92;
bill 20 mm.

Nos. 2303, 2325, 2326, 2335, 2339. 9 9 ad. &imm. Névita:
17, 23, 25, 26.x1.08—Wing 135-140; tail 84-87; bill 18-
19 mm.

No. 2274. (¢) juv. Novita: 12.x1.08.—Wing 154; tail 100;
bill 18 mm.

‘Tris orange (or orange-yellow), feet and bill black.”

“J have seen, but been unable to shoot this bird in all parts
of the Chocé from Malaquita (S. T.) up to Sipi” (M. G. Palmer).

The series of males illustrates beautifully the variation accord-
ing toage. In perfectly adult birds the six outer primaries and
the central pair of rectrices bear only a small, rounded black spot
at the tip, while the plumage otherwise is pure white, some-
times with a scarcely perceptible greyish tinge on the pileum.
Immature males have much more black on the wings, and most,
if not all, of the rectrices tipped with black. The younger the
bird, the more extended is the black colour. No. 2287 has not
only the black tips to the three outer primaries much larger than
tht specimens in more advanced stage, but the greater part of the
' inner web of the remaining primaries isalso black. The shafts of
the primaries are pure white in adult, black in immature males.
The latter, besides, have the bastard-quills and primary coverts
edged or tipped with black.

No. 2273 is changing from the grey juvenile plumage into the
white of the adult. Another, still younger, male, No. 2274,
resembles the females, but may be distinguished by its larger size,
much paler cinereous upper parts and breast, whiter belly, ete.
A specimen in the Tring Museum from Ventana, N.W. Ecuador,
is exactly like it.

C. hopket is peculiar to the hot, forest-covered lowlands of

1148 MR. C. E. HELLMAYR ON THE

Western Colombia and the adj joining parts of Ecuador (province
Esmeraldas).

101. QuERuLA puRPURATA P. L. S. Mull.

Muscicapa purpurata P. lL. 8. Miller, Natursyst. Suppl.
p. 169 (1776—ex Daubenton, Pl. Enl. 381 : Cayenne).

Querula cruenta Cassin, Proc. Acad. Philad. 1860, p. 143
(Turbo); Sclater & Salvin, IPS a Se USD, jon" BAO (Pocune,
Remedios).

Nos. 2001, 2151, 2265. ¢ gd ad. Noanama: 22.vii.; Sipi:
3.x.; Névita: 12.xi.08.—Wing 178-182; tail 114-117; bill
24 mm. ;

Nos. 2044, 2152, 2266. 9 2 Noanama: J.ix.; Sipi: 3.x. ;
Novita: 12.x1.08.—Wing 166-168; tail 108-112; bill 22—
233 mm.

* Tris dark brown, feet black, bill blue.”

These birds have generally smaller bills than those from
Guiana, Venezuela (Caura) and Parad, but the difference is not
quite constant. ‘This is a wide- spread species, ranging from
Costa Rica to Peru, and the mouth of the Amazons.

102. SYNALLAXIS PUDICA Scl.

Synallaxis pudica Sclater, P. Z. 8. 1859, p. 191 (1859.—
Bogota) ; Sclater & Salvin, P. Z 8. 1879, p.-521 (Remedios,
Antioquia).

No. 2116. 9 ad: Sipi: 26.1x.08—Wing 60; tail 73; bill
15 mm.

No. 2730. 2 juv. Pueblo Rico, 5200 ft.: 16.viii.09.

“ Tris brown, feet blue-grey (no. 2116), dark brown (no. 2730),
bill black, mandible grey.”

Identical with specimens from Bogota collections.

103. SYNALLAXIS UNIRUFA Lafr.

Synnalaxis (sic) unirufa Lafresnaye, Rev. Zool. vi. p. 290
(1843.—* Colombia ”).

Synallaxis unirufa Sclater & Salvin, P. Z.S. 1879, p. 521
(Antioquia).

No. —. g ad. Tatamaé Mountain, OU ft., 9.x.09.—Wing
61; tail 70 mm. ; bill damaged.

«i Iris brown, lees dark grey, bill black.”

Agreeing with a Bogotdé skin. [he plumage is uniform clear
chestnut rufous, vather paler underneath, except, of course, the
blackish inner webs of the remiges. There is no black cular
patch, so characteristic a feature in the allied S. castanea Sel. iy
from the Silla de Caraccas, Venezuela.

S. unirufa is peculiar to the high mountains of Colombia.

* Ann. Mag. Nat. Hist. (2) xvii. p. 466 (1856.—Caraccas).

BIRDS OF WESTERN COLOMBIA. 1149

104. SIproRNIs ERYTHROPS GRISEIGULARIS Ridgw.

[Synallaxis erythrops Sclater, P. Z. S. 1860, p. 66 (1860.—
Pallatanga, W. Ecuador). |

Acrorchilus erythrops griseigularis Ridgway, Proc. Biol. Soc.
Wash. xxi. p. 72 (1909.—San Antonio, Rio Cali, ““N.W.”
Colombia; M. G. Palmer coll.).

Synallaxis erythrops (nec Sclater) Sclater & Salvin, P. Z.S8. 1879,
p. 521 (Frontino, W. Cordillera).

Nos. 3740, —. 6 ¢ ad. Pueblo Rico (5200 ft.), 7.1x.; Siat6
(5200 ft.), ix.09.— Wing 69; tail 63, 65; bill 13 mm:

No. 2799. 2 juv. Loma Hermosa, R. Jamaraya (4150 ft.),
18.x.09.— Wing 63: tail 68; bill 13 mm.

“ Tris brown, feet greyish green, maxilla black, mandible light
brown.”

The adult birds fully bear out the characters given by Mr.
Ridgway for his subspecies which, moreover, was based upon
one of Palmer’s skins. SS. ¢. griseigularis may easily be recog-
nized from its allies: S. e. erythrops Scl., of Western Heuador,
and 8. e. rufigenis Lawr.*, of Chiriqui and Costa Rica. by its
mouse-grey (instead of buffy brown) foreneck and breast, and
brighter cinnamon-rufous median rectrices.

S. e. griseigularis is peculiar to the elevated districts of the
Western Cordillera of Colombia.

[105. THriPADECTES SCLATERI Berl.

Thripadectes sclateri Berlepsch, Ornis, xiv. p. 365 (Feb. 1907.—
S. Pablo, W. Colombia, 4500 ft. ; G. Hopke coll.).

Rhopoctites alogus Bangs, Proc. Biol. Soc. Wash. xxiii. p. 72
(1910.—Pavas, W. Colombia, 4400 ft.; M.G. Palmer coll.).

Count Berlepsch having kindly submitted to my inspection the
type specimen, 1 can positively state that f. alogus is merely a
synonym of 7’. sclateri, the description of which appears to have
been overlooked by Mr. Bangs. The two types were obtained in
the same district, viz., in the Western Cordillera above Buena-
ventura, Chocé Bay. Although somewhat resembling Automolus
(Rhopoctites) rufobrunneus in general coloration, 7’. sclateri isa very
distinct species, and I agree with its describer that it finds its
natural place in the genus Thripadectes. The structural characters,
especially the shape and size of the bill, are exactly the same as
in 7. flammulatus, while A. rufobrunneus has a much shorter,
slenderer, more depressed bill, and much shorter, weaker tarsi.
The pileum has distinct, though narrow, pale shaft-lines, whereas
it is uniform dusky olive in A. rufobrunneus ; the back is darker,
less reddish ; the throat much paler, light ochraceous instead of
deep orange; the sides of the neck are olive-brown instead of
orange-ochraceous ; the breast and abdomen are darker ochreous

* Synallaxis rufigenis Lawrence, Ann. Lyc. N. H. N.Y. ix. p. 105 (1868.— Costa
Riva).

1150 MR. GC. E. HELLMAYR ON THE

and lack the pale shaft-stripes, so conspicuous in A. rufo-
brunneus, ete. etc.

The type, an adult female, obtained by the late Gustav
Hopke, March 6, 1897, measures—wing 99; tail 95; bill
27 mm. |

106. AUTOMOLUS NIGRICAUDA Hart,

Automolus nigricauda Hartert, Bull. B. O. C. vil. p. xxx,
(1898.—Cachabi, N.W. Ecuador); idem, Nov. Zool. v. 1898,
p. 491 (Cachabi); idem, 1. ¢. ix. 1902, p. 616 (Paramba, 3500 ft. ;
Rio Sapayo, 450 ft.: N.W. Ecuador).

No. 2414. g ad. Noévita, 150 ft., 18.x11.08.— Wing 83 ; tail 71;
bill 23 mm.

“ Tris light brown, feet and bill black.”

This bird is practically identical with the specimens in the
Tring Museum. ‘The species was hitherto known only from
N.W. Ecuador, where Mr. Rosenberg and his collectors obtained
three examples at Cachabi (500 ft.), Paramba (3500 ft.), and Rio
Sapayo (450 ft.).

A. nigricauda is nearly allied to the group of A. rubiginosus
from Central America, but may easily be distinguished by its dull
blackish (instead of bright chestnut rufous) tail, less rufous wings,
darker ferruginous foreneck, dull brownish olive belly, etc., ete.

107. HytoctistEs SUBULATUS ASSIMILIS Ber]. & Tacz.

[Sphenera subulata Spix, Av. Bras. i. p. 82, pl. Ixxxii. fig. 1
(1824—‘* in sylvis fl. Amazonum ”). |

Automolus assimilis Berlepsch & Taczanowski, P. Z. 8S. 1883,
p. 561 (1883.—Chimbo, 8.W. Ecuador); Hartert, Nov. Zool. v.
1898, p. 491 (Chimbo, Cachabi, W. Ecuador).

No. 2080. $ (2) ad. Sipi, Rio Sipi: 12.ix.08.—Wing 82;
tail 71; bill 203 mm.

No. 2467. 3 (%) ad. Noanama: 13.1.09.—Wing 80; tail 69;
bill 20 mm.

Nos. 2725, 2701. 6 Qad. Tadé: 8, 28.vi.09.—Wing 87, 86;
tail 70; bill 21, 22 mm.

No. 2093. @ ad. Sipi: 19.ix.08—Wing 83; tail 67; bill
20 mm.

“ Tris dark brown, feet greyish green or black brown, maxilla
black, mandible light brown.”

This series agrees perfectly with two specimens from Western
Keuador (Carondelet, Paramba) in the Munich Museum, The
upper part of the head is nearly uniform olive-brown with
obsolete dusky edges to the feathers; the back wholly unstriped,
rufescent brown; the upper wing-coverts and outer webs of the
remiges are deep russet-brown; the breast and abdomen dull
light olivaceous, the chest but indistinctly mottled with buffy,
etc. etc.

BIRDS OF WESTERN COLOMBIA, ILI

The Costa Rica form, H. s. virgatus Lawr.*, may be distin-
guished from H. s. assinvilis by its more elongated as well as
slenderer bill (234 to 24 mm.); bright chestnut-brown colour
of the wing-coverts and remiges; blackish ground-colour of the
pileum and nape, with sharply defined, buff shaft-streaks; by
having the upper back distinctly streaked with buff, ete.

Mr. Ridgway has rightly separated P. virgatus under the new
generic term Hyloctistes ~, but it is certainly only subspecifically
distinct from H. s. subulatus and H. s. assimilis, the three races
replacing each other geographically. With regard to the differ-
ences between the two last-named races see my remarks in Noy.
Zool. xvii. 1910, p. 320.

108. XENICOPSIS SUBALARIS SUBALARIS Scl.

Anabates subalaris Sclater, P. Z. S. 1859, p. 141 (1859.
tanga, W. Ecuador).

Anabazenops subalaris Sclater, Cat. B. Brit. Mus. xv. p. 108
(part. : a-c, Pallatanga; d, Quito).

No.—. ¢ ad. Loma Hermosa, R. Jamaraya, 4150 ft.,
19.x.09.— Wing 91; tail 84; bill 19 mm.

“Tris dark grey, feet grey-green, maxilla black, mandible
yellow.”

This bird is somewhat difficult to place. In coloration, notably
in the dark brown ground-colour of the pileum and back, it agrees
with topotypical West Ecuadorian specimens; but it is fully as
large as the Central American form, X. s. lineatws Lawr. f,
which, however, has the head above and the back of a distinctly
paler, more olivaceous brown tinge. Until more specimens from
Western Colombia come to hand its identification can be regarded
only as provisional.

X. s. subalaris inhabits the elevated districts of the Western
Cordillera in Ecuador and Colombia, extending down the slope to
about 2000 feet.

X. mentalis Tacz. & Berl.§, united by Dr. Sclater to 1. s.
subalaris, is a very distinct species, though it may eventually
prove to be the Hastern representative. It differs particularly in
the following characters: the top of the head is much darker,
blackish, and covered with broad, buff shaft-stripes, while in
A. s. subalaris there are but a few narrow hair-like streaks to be
seen ; the upper back also shows much broader as well as more
numerous buff stripes; the whole under surface, posterior to
foreneck, is likewise broadly striped all over with yellowish

Paila-

* Philydor virgatus Lawrence, Ann. Lyc. N. H. N.Y. viii. p. 468 (1867.—Ango-
stura. Costa Rica).

+ Proc. Biol. Soc. Wash. xxii. p. 72 (1909.—Type: Philydor virgatus Lawr.).

t Anabazenops lineatus Lawrence, Ann. Lyc. N. H: N.Y. viii. p. 127 (1865.—
ncoctarss Costa Rica).

§ Anabazenops mentalis Taczanowski & Berlepsch, P. Z.S. 1885, p. 96 (1885.—
Machay, Eastern Ecuador).

1152 MR. C. E. HELLMAYR ON THE

white, etc., etc. The pattern of the upper parts reminds one
rather of Automolus holostictus Scl. & Salv., yet it need not be
emphasized that this species is generically distinct.

Of X. mentalis, I have examined two adult males and an
immature female from Machay, E. Ecuador, and two Bogota
skins, one in the British Museum, the other in Count Berlepsch’s
collection.

109. XENOPS GENIBARBIS LITTORALIS Scl.

[Xenops genibarbis Mliger, Prodr. Syst. Mamm. et Av. p. 213
(1811.—Cameta, Lower Amazons). |

Xenops littoralis Sclater, P. Z. S. 1861, p. 379 (1861.—Esme-
raldas, N.W. Ecuador).

Xenops ruficauda (nec Vieillot) Cassin, Proc. Ac. N, Sci. Philad.
1860, p. 193 (Turbo, N. Colombia).

Xenops genibarbis (nec Lliger) Sclater & Salvin, P. Z.8. 1879,
p. 523 (Remedios).

No. 2039: ¢ imm. Noanama: 31.viii.08.—Wing 61; tail 46;
bill 123 mm.

No. 2479. 9 ad. Noanama: 15.1.09.—Wing 61; tail 463;
bill 15 mm.

No. 2646. g ad. Tadé, R. San Juan: 17.v.09.—Wing 623;
tail 48 ; bill 13 mm.

“ Tris dark brown, feet blue-grey, bill black.”

These birds agree with topotypical examples from the Ecuadorian
province of Esmeraldas. _X. g. mexicanus Scl.*, which ranges
from E. Mexico to Panama, is more rufescent, less olive, under-
neath, and has the back of a brighter, clearer cinnamomeous-
brown.

X. q. littoralis inhabits the lowlands and foothills of Western
Eeuador and Western Colombia.

110. GuyPHORHYNCHUS CUNEATUS CASTELNAUDIT Des Murs.

[Dendrocolaptes cuneatus Lichtenstein, Abhand], Akad. Berlin
a.d. Jahren 1818-19, p. 204, pl. un. fig. 2 (1820—“‘in Brasilice
provincia Bahia”; cfr. idem, |. c. a.d. Jahren 1820-21, p. 264. |

Glyphorhynchus castelnaudii Des Murs, in Castelnau’s Voyage,
Oiseaux, p. 47, pl. xv. fig. 2 (1856.—Santa Maria, Hastern Peru).

Glyphorhynchus cuneatus (evrore) Sclater & Salvin, P. Z.8. 1879,
p- 523 (Remedios).

Nos: 2019 2d: o mm. Orad.) peiprs) L0set Neamamanr
26.vili.08.— Wing 69, 67; tail 70, 62; bill 12 mm.

“Tris dark brown, feet dark grey, maxilla black, mandible

rey.”

Slightly smaller than Upper Amazonian skins, but similar in

coloration.

_ %* Xenops “ mexicanus Sclater, P. Z. S. 1856, p. 289 (Jan. 1857.—Cordova;
E. Mexico).

BIRDS OF WESTERN COLOMBIA. 1158

111. DENDRORNIS LACHRYMOSA ROSTRATA Ridew.

[Dendrornis lachrymosus Lawrence, Ann. Lye. N. ET INE ONES ‘val
p. 467 (1862.— Panama). |

Xiphorhynchus lacrymosus rostratus Ridgway, Proc. Biol.
Soc. Wash. xxii. p. 73 (1909:—Rio Dagua, ‘¢ North-western ”
Colombia).

Dendrornis lacrymosa (errore) Sclater & Salvin, P. Z.8. 1879,
p. 523 (Remedios) ; Hartert, Nov. Zool. ix. 1902, p. 616 (N.W.
Keuador).

Dendrornis — sp. Cassin, Proc. Acad. N. Sci. Philad. 1860,
p. 194 (R. Truando).

Nos. 2587, 2615. g¢ gad. Condoto, R. Condoto, NBO) siaey 3
6,20.iv.09.—Wing 126, 122; tail 103, 98; bill 837 mm.

Nos. 2004, 2005. dg imm., 2 imm. Noanama, 1000 fia: 24,
26.viii.08.— Wing 126, 120; tail 105, 102; bill 35, 32 mm.

“Tris dark brown, feet green, maxilla black, mandible grey.”

These specimens, as well as several others from North-western
Ecuador (Bultin, Carondelet), have the bill slightly deeper and
stouter than typical birds from Panama. The culmen, how-
ever, is not more strongly arched. I must confess I am not
very confident as to the distinctness of this race and should not
be surprised if a larger series would show it to be inseparable
from D. 1. lachrymosa. D.1. rostrata, if really distinguishable,
ranges from N.W. Colombia (Rio Truando) along the Pacific coast
southwards to the north-western portion of Ecuador. It is found
only in the lowlands and on the hill-slopes.

112. DENDRORNIS TRIANGULARIS ZQUATORIALIS Berl. & Tacz.

[Dendrocolaptes triangularis Latresnaye, Rev. Zool. v. p. 134
(1842.—“ Bolivia,” * errore; the type came from Santa-Fé-de-
Bogota, Colombia). |

Dendrornis erythropygia equatorialis Berlepsch & Taczanowski,
P. Z. 8. 1883, p. 563 (1883.— Chimbo, W. Keuador) ; Hartert,
Nov. Zool. v. 1898; p. 490 (Chimbo, Paramba, W. Eeuador).

D. cequatorialis Salvadori & Festa, Boll. Mus. Torino, XIV.
no. 362, 1899, p. 25 (R. Peripa, W. Ecuador).

D. punctigula (errore) Goodfellow, Ibis, 1902, p. 63 (Nanegal,
Intac, Gualea, W. Ecuador; specimens examined).

No. 2378. g ad. Névita: 7.x11.08—Wing 117; tail 103;
bill 34 mm.

* The locality “ Bolivia” is certainly a mistake, for in Mag. Zool. 1843, Oiseaux,
pl. 32, Lafresnaye expressly says: ““ vient de Santa-Fé-de-Bogoté.” This statement
is repeated in Rev. Mag. Zool. (2) ii. 1850, p. 419, in the French text, while the
Latin diagnosis (a verbatim reprint of the original account) terminates with the
indication: “habitat in Bolivia.” Moreover, at that time, Bogoté skins reached
Paris in large numbers, whereas material from Bolivia was very scarce in European
Museums. In fact, D'Orbigny was the only naturalist to send collections from that
country, but he did not obtain the species in question.

1154 MR. CG. E. HELLMAYR ON TIE

No. 2529. gad. El Tigre, 320 ft.: 10.11.09.—Wing 116;
tail 103; bill 834 mm.

“Tris dark brown, feet grey-green or blue, maxilla black, tip
light brown, mandible grey.”

The two skins agree perfectly with a series from Western
Ecuador in the Munich Museum. JD. ¢. cequatorialis is an
excellent form, differing in many important details from the
typical race of Central Colombia, W. Venezuela, etc. The rump
and the wings are much darker chestnut rufous; the ground-
colour of the throat is decidedly buff, instead of whitish ; the dark
olive markings on the throat are restricted to small, rounded
apical spots, while in the allied D. ¢. triangularis the feathers
show a continuous, marginal edge, giving a squamate appearance ;
the top of the head is wholly unstreaked, plain dark olive, only
sometimes with a few buff shaft-lines on the forehead, whereas
the typical form has the entire crown conspicuously streaked or
spotted with pale buff, ete.

D. t. punctigula Ridgw. * from Costa Rica and Chiriqui, is
most closely related to D. t. equatorialis, agreeing in the pattern
of the throat and in the uniform pileum, but differs in the clearer
rufous rump and wings, the deeper olive-green ground-colour, and
the smaller, less rounded spots of the lower parts. Single
examples are, however, not always distinguishable.

D. t. triangularis shows a distinct, whitish lateral stripe near
the lower edge of the maxilla, while in D. t. equatorialis and
D. t. punctigula the upper bill is uniform dark horn-brown or
blackish. This difference is quite constant in the enormous series
which I have examined.

D. ¢. equatorialis is strictly confined to the western slopes of
the Andes in Ecuador and Southern Colombia. In Ecuador it
ranges from sea-level up toabout 3500 feet, for we have specimens
from Ventana (90 feet) and Bultn (160 feet) as well as from Lita
(3200 feet) and Paramba (3500 feet). From Colombia, west of
the Coast Cordillera, I have seen only Palmer’s two skins, one
taken at 150, the other at 320 feet elevation 7.

D. t. triangularis inhabits the Central and Eastern Cordillera
of Colombia, the mountains of Western Venezuela (Merida,
Cumbre de Valencia), the eastern slopes of the Ecuadorian Andes,
and Northern Peru. ‘Two specimens from the Cauca slope of the
Coast Cordillera—one, from Santa Hlena, in the British Museum,
the other, from Riolima, at Tring—also belong to this form, being
in every respect typical. Birds from Hastern HEeuador (Machay,

* Dendrornis punctigula Ridgway, Proc. U.S. Mus. xi. p. 544 (1889.—Naranjo,
Costa Rica).

+ My material of D. ¢. equatorialis consisted of the following specimens :—
6 from Chimbo, including the type (Mus. H.v. Berlepsch & Tring); 2 from Cachabi
(Tring); 7 Paramba, 6 Lita, 1 Ventana, 5 Bultin (Mus. Tring, Munich, Vienna, etc.) ;
3 Nanegal, 2 Nono, 1 Intag (Goodfellow coll—TIring), 2 Chocéd, W. Colombia
(Munich): 385 examples in all.

BIRDS OF WESTERN COLOMBIA. 1155

Baeza *) and North Peru (Chachapoyas) are absolutely identical
with those from Bogoté and Venezuela 7.

Ona future occasion [I hope to give some information respecting
several other species of the genus Dendrornis which, through
lack of proper material, are very imperfectly known.

113. DENDROCINCLA MERULOIDES LAFRESNAYEL Ridgw.

| Dendrosops meruloides Lafresnaye, Rev. Mag. Zool. (2) iii.
p. 467 (1851.—‘“ Cote ferme,” sc. Cumana, Venezuela). |

Dendrocincla lafresnaye. Ridgway, Proc. U.S. Mus. x. 1887,
p. 492 (1888.—‘“ Upper Amazons ?”—errore; we substitute
Colombia as type locality).

D. olivacea latresnayei Oberholser, Proc. Acad. N. Sci. Philad.
1904, p. 457 (crit.).

No. 2460. 9 (?) ad. Noanama: 11.1.09.—Wing 102; tail
88; bill 26 mm.

No. 2113. 9 imm. Sipi: 23.1x.08.—Wing 95; tail 81; bill
25 mm.

‘“‘ Tris dark brown, feet and bill black.”

These specimens agree well with others from Western Ecuador,
and Riolima, Cauca Valley. Bogota skins are slightly more
rufescent both above and below, thus pointing towards JD. J.
pheochroa Berl. & Hart.

D. m. lafresnayert, D. m. pheochroa, and D, m. meruloides are
clearly races of one and the same species, representing each other
geographically. The first-named form may always readily be
distinguished from the two others in having the outer webs of the
quills conspicuously washed with olivaceous, and the dusky
portions of the bill much darker, black instead of horn-coloured.
On the other hand, D. m. meruloides differs from D. i pheochroa
in its much brighter, more cinnamon or ochraceous-brown
coloration.

Range and principal synonyms of these forms are as follows :—

(a) D. MERULOIDES MERULOIDES Lafr.

Dendrocops meruloides Lafresnaye, Rev. Mag. Zool. (2) iii.
p- 467 [1851.—“ Cote ferme,” sc. Cumand; cfr. Ménégaux
& Hellmayr, Mém. Soc. Hist. Nat. Autun, xix. 1906, p. 121
crit.) |.

: Dee meruloides Cabanis & Heine, Mus. Hein. ii. 1859,
p. 34 (Caraccas); Sclater & Salvin, P. Z.8. 1868, p. 167 (Pilar,
Venezuela); Chapman, Bull. Amer. Mus. N.Y. vi. 1894, p. 48
(Trinidad) ; Berlepsch & Hartert, Nov. Zool. ix. 1902, p. 67

* D. erythropygia(!) Goodfellow, Ibis, 1902, p. 63 (Baeza, EH. Ecuador).

+ My material of D. t. triangularis was composed as follows :—9 Bogota (Tring,
Munich, H.& B.), 1 Santa Elena, 1 Riolima,Cauca (Brit. Mus., Tring), 10 Merida
(fring & Munich), 3 Cumbre de Valencia (Munich), 4 East Ecuador (2 Machay,
H. & B.; 2 Baeza, Tring), 2 N. Peru, Chachapoyas (Tring) ; 30 examples in all.

{ =D. olivacea auct. ex parte: America merid.

1156 MR. C. E. HELLMAYR ON THE

(Cumand, San Hsteban) ; Hellmayr, Nov. Zool. xiii. 1906, p. 29
(Trinidad).

D. meruloides aphanta Oberholser, Proc. Acad. N. Sei, Philad.
1904, p. 460 (Lobago).

Type locality : Cumana, Venezuela.

Hab. Tobago ; Trinidad ; North Coast of Venezuela: from
Cumanid, to Puerto Cabello.

Examined: 16 Trinidad, 2 Tobago, 6 Cumana, 1 Caraccas,
12 San Esteban (near Puerto Cabello).

(b) D. meruLorEs PHmocHROA Berl. & Hart.

Dendrocincla® pheochroa Berlepsch & Hartert, Nov. Zool. 1x.
p. 67 (1902.—Munduapo (type), Maipures, Nericagua, Orvinoco ;
Suapure, La Pricion, Caura R.) ; Thering, Rev. Mus. Paul. vi.
1905, p. 437 (Rio Jurud); Snethlage, Jou. f. Orn. 1908, p. 15
(Rio Purts).

D. olivacea pheochroa Oberholser, Proc. Acad. N. Sci. Philad.
1904, p. 458 (Suapure, La Union, Caura).

D. fumigata (errore) Pelzeln, Zur Orn. Bras. 1. 1867, p. 42
(part. Rio Branco, spec. examined) ; Taczanowski, P.Z.S. 1882,
p- 27; idem, Orn. Pérou, il. 1884, p. 168 (Huambo, N. Peru).

D. olivacea (nee Lawrence) Sclater, Cat. B. xv. p. 166 (part. :
I, m, Sarayacu, E. Ecuador; », Rio Napo ; q, Iquitos, INGuletere: 2
spec. examined),

Dendrocolaptes atrirostris (nec Lafr. & D’Orb.) Selater, P. ZS,
1858, p. 63 (Rio Napo; spec. examined).

Type locality: Munduapo, Orinoco R.

Hab. Amazonian region: from the Rio Branco, N. Brazil, and
the Caura Valley, Eastern Venezuela, westwards to N. Peru and
the eastern slopes of the Colombian and Ecuadorian Andes.

Examined : 7 Munduapo (inel. type), 2 Maipures, 2 Nericagua,
6 Caura, 1 Rio Branco, 1 R. Jurua, 2 N. Peru (Iquitos, R. Tigre),
3 Rio Napo, E. Ecuador, 3 Bogotsa.

(c) D. MERULOIDES LAFRESNAYEI Ridgw.

D. lafresnayei Ridgway, Proc. U.S. Mus. x. 1887, p. 492
(1888.—“ Upper Amazons,” errore).

D. olivacea lafresnayei Allen, Bull. Amer. Mus. xii. 1900, p. 156
(Santa Marta); Oberholser, Proc. Acad. N. Sci. Philad. 1904,
p. 457 (crit.).

Dendrocops atrirostris (errore) Sclater, P.Z.S. 1860, p. 66
(Pallatanga, W. Ecuador) ; idem, l.c. p. 278 (Babahoyo) ; idem,
lc. p. 293 (Esmeraldas).

Dendrocincla atrirostris Taczanowski, P. Z.8. 1877, p. 332
(Palmal, 8.W. Heuador); Berlepsch & Taczanowski, P. Z.8. 1883,
p. 563 (Chimbo, 8.W. Eeuador).

D. olivacea Sclater, Cat. B. Brit. Mus. xv. p. 166 (part. :
Colombia (Bogoti, Manaure), W. Ecuador); Hartert, Nov. Zool.

* Spelt Dendrocinda.

BIRDS OF WESTERN COLOMBIA. eae

v. 1898, p. 491 (Paramba, N.W. Ecuador); Goodfellow, Ibis, 1902,
p- 63 (Sto. Domingo, W. Ecuador); Salvadori & Festa, Boll. Mus,
Zool. Torino, xiv. no. 362, 1899, p. 27 (Vinces, W. Ecuador).

D. 0. anguina Bangs, Proc. Biol. Soc. Wash. xx. p. 138
(1898.—Santa Marta); idem, l.c. xi. 1899, p. 100 (Sierra
Nevada de Santa Marta).

Hab. Colombia (Bogoti, Santa Marta, Riolima, Chocé, ete.) and
Western Ecuador (Paramba, 8. Javier, Esmeraldas, Pallatanga,
Babahoyo, Vinces, Chimbo, Santo Domingo, Palmal, etc.), from
sea-level up to 6000 feet.

Examined: 15 Western Ecuador, 1 Santa Marta (Manaure),
1 Riolima, Cauca, 2 Chocé, 5 Bogota.

N.B.—The original locality ‘‘ Upper Amazons” is doubtless
erroneous, the species being strictly confined to the western slopes
of the Andes. In the Amazonian district it is replaced by the
preceding form.

114. CYMBILANIUS LINEATUS FASCIATUS Ridgw.

|Lanius lineatus Leach, Zool. Misc. i. p. 20, pl. vi. (1814.—
Berbice, British Guiana). |

Cymbilanius lineatus fasciatus Ridgway, Proc. U.S. Mus. vi.
1883, p. 415 (1884.—Rio Sucio, Costa Rica).

C. lineatus (nee Leach) Sclater & Salvin, P. Z.8. 1879, p. 524
(Remedios, Neche).

No. 2571. ¢ imm. Condoto: 27.11.09.—Wing 72; tail 69;
bill 21 mm.

No. 2572. Q ad. Condoto: 27.11.09.—Wing 72; tail 68;
bill 21 mm.

“ Tris red-brown, feet blue-grey, maxilla black, mandible grey.”

Specimens from W. Colombia and W. Ecuador have the black
barring of the lower parts slightly broader than those from
Cayenne and Amazonia. Although topotypical Costa Rican
birds are not available for comparison, | think they belong to
Ridgway’s subspecies.

Cl. fasciatus is confined to the lowlands and foot-hills.

115. 'THAMNOPHILUS MAJOR TRANSANDEANUS Sel.
[Thamnophilus major Vieillot, Nouv. Dict. 11. p. 313 (1816.—

ex Azara: Paraguay). |

T. transandeanus Sclater, P. Z. 8. 1855, p. 18 (1855.—Guay-
aquil, S.W. Ecuador) ; Cassin, Proc. Acad. N. Sci. Philad. 1860,
p- 188 (Ttirbo).

No. 1972. g ad. Guineo, Rio Calima, 6.vii.08.—Wing 92;
tail 70; bill 27 mm.

No. 2528. 9 ad. El Tigre, R. Tamana, 320 ft., 10.11.09.—
Wing 91; tail 69; bill 27 mm.

No. 3777. g ad. la Selva, R. Jamaraya, 4600 ft., 1.x.09.—
Wing 94; tail 70; bill 28 mm.

‘“‘ Tris vermilion, feet blue, bill black.”
Proc. Zoou. Soc.—1911, No. LX XVIII. 78

1158 MR. C. E. HELLMAYR ON THE

The Tring Museum possesses an adult male obtained by
Mr. Rosenberg at Rio Dagua (Juntas), May 27,1895. The males
from W. Colombia agree, in coloration and size, with a series from
Western Ecuador and Chiriqui, the under tail-coverts being
black with distinct white apical margins.

T. m. transandeanus is, otherwise, known only as an inhabitant
of the lowlands (from sea-level up to 1000 feet), and its occurrence
at La Selva (4600 feet alt.) must be regarded as exceptional.

In the Central and Eastern Cordilleras it is replaced by a nearly
allied, but easily recognizable race, 7. m. granadensis Cab. ™*,
which forms the passage to 7. m. melanurus Gould, of Upper
Amazonia. It differs, in the male sex, from 7. m. transandeanus
in having the under tail-coverts cinereous with a white apical
edge preceded by a distinct blackish subterminal band, and in
its much slenderer, weaker bill. I have examined several Bogota
skins in the collections of the Paris and Munich Museums, an
adult male from Antioquia (Salmon), and one from near
Mérida (Briceno) at Paris, and four males from N.W. Venezuela
(3d ad, g¢ imm., Mt. Bucarito, Tocuyo; ¢ ad. Hjido, near
Mérida; ¢ ad. San Esteban—Mocquerys coll.) in the Tring
Museum. All of them very clearly show the characters indicated
above. Cfr. also Ménégaux & Hellmayr, Bull. Soc. Philom.
Paris (9) viii. 1906, pp. 25-26.

116. THAMNOPHILUS NEVIUS ATRINUCHA Salv. & Godm.

[Lanius nevius Gmelin, Syst. Nat. 1, 1. p. 308 (1788—
ex Latham : Cayenne). |

Thamnophilus atrinucha Salvin & Godman, Biol. Centr.-Amer.,
Aves, ii. p. 200 (1892.—“‘ Central America,” no type specified ; we
fix Panama as terra typica).

T. nevius (nec Gmelin) Sclater & Salvin, P. Z.S. 1879, p. 524
(Neche, Antioquia); Berlepsch, Journ. f. Orn. 1884, p. 307
(Bucaramanga); Cassin, Proc. Acad. N. Sci. Philad. 1860, p. 188
(Turbo).

No. 1967. ¢ imm. San Joaquim: 4.viii.08.—Wing 71; tail
61; bill 184 mm.

No. 2049. ¢ imm. Noanama: 2.1x.08.—Wing 70; tail 56;
bill 18 mm.

No. 2127. gad. Sipi: 28.ix.08.—Wing 70; tail 54; bill
183 mm.

Nos. 2411, 2425, 2435. § gd ad. Ndéovita: 18, 22, 24.xii.08.—
Wing 70-74; tail 56-59; bill 183 mm.

Nos. 2395, 2428, 2431, 2436. 9 2. No6vita: 12,23, 24,
26.xii.08.— Wing 68-69 ; tail 54-58; bill 17-183 mm.

‘“‘ Tris brown or red-brown in male, grey in female, feet blue or
blue-grey, maxilla black, mandible grey.”

Specimens from Colombia and Western Ecuador are practically

* Diallactes granadensis Cabanis, Journ. f. Orn. 1872, p. 234 (Bogota coll.).

+ 7. transandeanus (errore) Sclater & Salvin, P.Z.S. 1879, p. 524 (Remedios,
Neche: Antioquia).

BIRDS OF WESTERN COLOMBIA. 1159

identical with others from Costa Rica, 7. . atrinucha replaces
7’. n. nevius on the western sides of the Andes, while the typical
form is found in the Guianas, the Orinoco Valley and _ its
tributaries.

For the distinguishing characters and geographical distribution
of the various races of this group see my remarks in Abhandl.
Bayer. Akad. Wissensch. II. Cl., vol. xxii. 3, 1906, p. 658 ff.

117. MyRMOTHERULA SURINAMENSIS PACIFICA, subsp. n.

No, 2132. gd ad. Sipi: 30.1x.08.—Wing 53; tail 30; bill
16 mm.

Nos. 1974, 2133. 9 9 ad. Guineo, R. Calima: 6.viii.08 ;
Sipi: 30.1x.08.— Wing 52, 50; tail 30; bill 143, 15 mm.

[No. 09.788, g ad. Buenaventura, 23.ii1.99—Wing 522;
tail 29; bill 15 mm.

No. 09.787. g ad. Plano de los Monos, near Naranjo, 2800 ft.,
3.iv.99.— Wing 53; tail 313; bill 15 mm.

No. 09.789-791. 9 9 ad. Buenaventura: 11,18, 23.iii1.99.—
Wing 51-52; tail 28-30; bill 144-15 mm.

These five specimens were collected by Hugéne André. |

“ Tris dark brown, feet blue-grey, bill black, mandible grey.”

Adult. Differs from J/. s. surinamensis Gm., in its longer
tail, much stronger as well as decidedly longer bill, wider white
tips to the outer rectrices (about 3 to 4 mm. on outermost pair),
and in having the upper back less variegated with black, while
the white interscapular blotch is much smaller in the male and
wholly absent in the female. The latter sex, too, has the top of
the head conspicuously clearer, about “tawny ochraceous” (Ridgw.
Nomencl. v. fig. 4) instead of “deep tawny ” (l.c. v. fig. 1), and
the dark stripes on the hind crown and nape are much duller and
less pronounced, being pale dusky olive instead of deep black.

Type in the Zoological Museum, Munich: no. 09.789. 9 ad.
Buenaventura, Chocd, W. Colombia, March 11, 1899. E, André
coll.

Besides the above, I have examined in the Tring Museum
three males and three females obtained at Juntas, Rio Dagua, by
Messrs. Rosenberg and Raap, and in the British Museum
a single female from Remedios, collected by T. K. Salmon.
Dr. Hartert * has already alluded to this form, and, after studying
large series from various localities, I have no hesitation in
separating the Pacific birds from true MZ, s. surinamensis.
The typical race is confined to Surinam, French and British
Guiana, extending westwards to the Caura Valley, Eastern
Venezuela, while JM. s. pacifica ranges from Western Ecuador
through Colombia north to Veragua. The females of both forms
have the sides of the head, the foreneck and chest uniform
ochraceous, thereby differing very conspicuously from the
Amazonian race J. s. multostriata Scl.t.

** Nov. Zool. ix. 1902, p. 612.
+ Cfr. Hellmayr, Noy. Zool. xvii, 1910, p. 345. :
78

1160 MR. C. E, HELLMAYR. ON THE,
Characters and range of the two races are as follows :—

(a) M. SURINAMENSIS SURINAMENSIS Gm.

Sitta surinamensis Gmelin, Syst. Nat. 1, i. p. 442 (1788—
based on: “ Surinam Nuthatch,” Latham, Gen. Syn. 1, 11. p. 654,
pl. 28 (= @ ): Surinam).

Formicivora quadrivittata (ex Lichtenstem MS.) Cabanis,
Arch. f. Naturg. 13, i. p. 227 [1847.—Guiana (Schomburgk)}.

Myrmotherula surinamensis Sclater, Cat. B. Brit. Mus. xv.
p. 231 (part.: m-g, Brit. Guiana; 2, ata ee Salvin, Ibis,
1885, p. 425 (Camacusa) ; Berlepsch & Hartert , Nov. Zool. ix.
1902, p. 73 (Caura); Berlepsch, Nov. Zool. xy. p. 154 (Cayenne).

Adult. Bill short and slender; white tips to external rectrices.
restricted, not more than 14 to 3 mm. wide on outermost pair.
Upper back, in both sexes, mostly black, variegated with narrow
white edges, base of feathers extensively white, forming a large,
concealed dorsal patch. Female with upper part of the head deep
tawny, crown and nape broadly streaked with deep black; sides
of the head, throat, foreneck and chest bright ochraceous, middle
of throat sometimes paler, ochraceous-buff or buff: rest of lower
parts buff, flanks shaded with greyish. On the sides of the neck
and chest there are sometimes a few faint dusky shaft-lines, but
in most specimens these parts are quite uniform.
Hab. Surinam: Paramaribo (Chunkoo: Tring Museum) ;
French Guiana: Approuague, Ipousin (Cherrie: Tring Museum),
Saint-Jean-du Maroni (Ze Moult: Munich Mus.) ; British
Guiana : Camacusa, Takutu R., Carimang R. (Whitely). Hast
Venezuela, Caura Valley: Nicare, La Pricion, Suapuré, Mato R.
(André, Klages: Tring and Munich Mus.) *
Material. 22 specimens. Birds from various localities measure
as follows :—
Two ¢ gad. Camacusa, British Guiana.— Wing 50, 52; tail 25,
26; bill 14, 15 mm.

Two ¢ ¢ ad. Paramaribo, Surinam.—Wing 50, 52; tail 25,
253; bill 14 mm.

Three ¢ g ad. French Guiana.—Wing 49-51; tail 24-25 ;
bill 14-15 mm.

Five ¢ gd ad. Caura R., Venezuela.—Wing 52; tail 25- a
bill 14-15 mm.

Two 9? 9 ad. Surinam.—Wing 49; tail 243-25; bill 14 mm.

One 2 ad. Carimang R.., Bulbine Ghote, —Wing 502; > tal 26%
bill 143 mm.

Two 9 9 ad, French Guiana.—Wing 49, 50; tail 24, 25 ;
bill 14 mm.

Five 9 9 ad. Caura R., Venezuela— Wing 49-50; tail 25—
27; bill 143-15 mm.

* Tt must remain doubtful, in the absence of females, whether the single male from

Munduapo, Upper Orinoco (ef. Nev. Zool. ix. p. 73) is referable to MW. s. surinamensis -
or to M. s. multostriata.

BIRDS OF WESTERN COLOMBIA, 1161

(6) M. suRINAMENSIS PaciFIcA Hellm.
WM. surinamensis (nee Gmelin) Cassin, Proc. Acad. N. Sci. Philad.
1860, p. 190 (R. Truando, N.Colombia); Lawrence, Ann. Lyc. N. H.
N.Y. vii. 1861, p. 293 (Isthmus of Panama); Sclater & Salvin,
P.Z.S. 1864, p. 356 (Lion Hill, Panama); Sclater & Salvin,
P.Z.S. 1879, p. 525 (Remedios); Berlepsch & Taczanowski,
P. Z.S. 1883, p. 564 (Chimbo, 8.W. Ecuador); Sclater, Cat. B.
Brit. Mus. xv. p. 231 (part.: a-g,j, k, 1); Salvadori & Festa,
Boll. Mus. Zool. Torino, xiv. no. 362, 1899, p. 29. (part.: Peripa,
W. Ecuador) ; Hartert, Nov. Zool. ix. 1902, p. 612° (San Javier,
Pambiltr, N.W. Ecuador); Goodfellow, Ibis, 1902, p. 64
(S. Nicolas, Gualea, W. Ecuador).
“ Myrmotherula ?” Sclater, P.Z.S. 1860, p. 294 (Esmeraldas,
N.W. Keuador).
Adult. Bill strong and heavy ; white tips to external rectrices
larger, varying from three to five millimetres in width on the
outermost pair. Upper back, in the male, much less variegated
with black, the white dorsal patch much less extended. Female
with upper part of the head much lighter, tawny ochraceous, crown
and hind neck with narrow, rather indistinct, dusky olive shaft-
streaks; upper back mainly pale grey, rather sparingly spotted
with black and edged with white, but without any trace of the
white dorsal blotch. Sides of the head and lower parts exactly as
in the female of MW. s. surinamensis.
Hab. Western Ecuador: Peripa (Festa), Intac (Buckley),
San Nicolas, Gualea (Goodfellow d& Hamilton), Chimbo (Siemi-
radzki), Esmeraldas (raser), San Javier, Pambilar (Plemming &
Miketta). Western Colombia: Buenaventura (André), Guineo,
Sipi (Palmer), Juntas (Rosenberg, Raap), Naranjo (André),
Remedios (Salmon), R. Truando (Wood). Isthmus of Panama:
Lion Hill (McLeannan), San Pablo (Salvin), Veragua (Arcé).
| Also in Bogoté coll. |
Material. 41 specimens. Birds from various localities average
as follows :—
Hight dg gd ad. W. Ecuador.—Wing 51-53; tail 28-30 ;
bill 16 mm.

Six ¢ dad. W. Colombia, Chocé district—Wing 523-54 ;
tail 29-314; bill 153-164 mm.

Two § 6 ad. Panama (Lion Hill)—Wing 51; tail 27, 28;
bill 153, 16 mm.

Thirteen 9 @ ad. W. Heuador.—Wing 49-52; tail 28-302 ;
bill 15-16 mm.

Hight ¢ 9. W. Colombia, Chocé district—Wing 50-52;
tail 28-303 ; bill 143-16 mm.

One @. Remedios, N.W. Colombia.—Wing 523; tail 304;
bill 15 mm.

-Two 2 Q@ ad. Bogoté.—Wing 52,53; tail 30,31; bill 15 mm.

One Q ad. Panama (Lion Hill)—Wing 50; tail 273; bill

15 mm.

1162 MR. C. E. HELLMAYR ON THE

118. MyrMornEeRULA FULVIVENTRIS VIDUATA Hart.

[Myrmotherula fulviventris Lawrence, Ann. Lyc. N. H. N.Y.
vil. p. 468 (1862.— Panama). |

Myrmotherula viduata Hartert, Nov. Zool. v. p. 492 (1898.—
Cachabi, N.W. Ecuador ; @ ).

M. fulviventris viduata idem, 1. c. ix. p. 612 (erit.).

M. ornata® Cassin, Proc. Acad. N. Sci. Philad. 1860, p. 191
(Rio Truando).

COE ?” Sclater, P. Z.S. 1860, p. 294 (Esmeraldas, N.W.
Keuador).

M. fulwiventris (nec Lawrence) Sclater & Salvin, P. Z.S. 1879,
p- 525 (Remedios, Antioquia) ; Wyatt, Ibis, 1871, p. 331
(Naranjo, near Bucaramanga); Salvadori & Festa, Boll. Mus.
Zool. Torino, xiv. no. 362, 1899, p. 29 (Peripa, W. Ecuador).

No. 2512. ¢ ad. El Tigre, 4.11.09.—Wing 53; tail 34;
bill 143 mm.

No.. 2426. ¢ juv. Noévita, 22.xi.08.—Wing 514; tail 38;
bill 14 mm.

No. 2471. 2 ad. Noanama, 13.1.09.—Wing 53; tail 38;
bill 143 mm.

“Tris dark brown, feet blackish, bill black, mandible grey.”

These specimens as well as several Bogota skins agree perfectly
with a series from W. Kcuador, including the type, kindly for-
warded by Dr. Hartert from the Tring Museum. JZ, f. viduata
differs from typical fulviventris in having the upper parts
decidedly more brownish or rufescent brown, this being especially
noticeable on the crown and upper tail-coverts; the edges to the
quills and rectrices are also more reddish, and the females have
the belly of a darker fulvous colour. J. f. fulviventris is known
to me from seven skins from Nicaragua (Rio Grande) and
Eastern Costa Rica (Carrillo, Reventazén, Matina R.), but Ihave
not yet seen any from the type locality.

M. f. viduata apparently ranges all over the Pacific lowlands,
up to about 2000 feet altitude, from the Rio Truando in the
north to the neighbourhood of Guayaquil, S.W. Ecuador, in
the south. There is an adult male in the Tring Museum from
Ana Maria (not far from Guayaquil), procured by Herr G. von
Buchwald in December 1905.

119. MyRMOTHERULA AXILLARIS MEZLNA Sel.

[Myrmothera axillaris Vieillot, Nouv. Dict. xii. p. 113 (1817.—
“ Guyane”).

Formicivora melena Sclater, P. Z. 8. 1857, p. 130 (Oct. 1857.—
Bogota).

Myrmotherula melena Sclater & Salvin, P. Z.S8. 1879, p. 525
(Neche, Antioquia); Cassin, Proc. Acad. N. Sci. Philad. 1860,
p- 191 (R. Truando).

No. 2416. ¢ ad. Névita: 19.xii.08.—Wing 53; tail 35;
bill 14 mm.

“Tris brown, feet and bill black.”

Agreeing with Bogota skins.

BIRDS OF WESTERN COLOMBIA. 1163

120. MyYRMOTHERULA SCHISTICOLOR SCHISTICOLOR Lawyr.

Formicivora schisticolor Lawrence, Ann. Lyc. N. H. N. Y. viii.
p- 173 (1865.—Turrialba, Costa Rica).

Myrmotherula schisticolor Hellmayr, Verhandl. zool.-bot. Ges.
Wien, lii. 1903, p. 210 (crit.).

No. 3765. 2. Siaté, Rio Siaté: 21.ix.08.—Wing 57; tail
39; bill 12 mm.

“Tris dark brown, feet plumbeous, bill black, mandible horn-
coloured.”

This bird agrees well with females from Costa Rica (Rio
Naranjo, Boruca), Chiriqui (Boquete), and Western Ecuador,
except in being slightly more greyish, less brownish, on the upper
parts. I have not seen male examples from W. Colombia, but
several from W. Ecuador, which I have before me, I am unable
to separate from typical Costa Rican skins.*

On the other hand, in the Sierra Nevada de Santa Marta,
N. Colombia, and in the mountains of Northern Venezuela
(Cumbre de Valencia, Cuman4), a well-marked geographical race,
M. schisticolor sancte-marte Allen, takes its place. The male
may be immediately recognized in having the black colour
restricted to the throat and middle of the foreneck, while in
M. s. schisticolor it extends over the breast down to the upper
abdomen. The females are not always distinguishable, though
as a rule those of sancte-marte have the back of a purer olive-
grey, without any brownish tinge. The Munich Museum
possesses a good series of this rare form from the Cumbre de
Valencia and the mountains of Cumana, Venezuela. The two
males from 8. Esteban, Venezuela, in the British Museum, Goering
coll.f, are likewise referable to W/. s. sancte-marte.

121. FoRMICIVORA QUIXENSIS CONSOBRINA Scl.

[| Thamnophilus quixensis Cornalia, Vertebr. Syn. Osculati Coll.
p. 12 (1849.—Quixos, Eastern Ecuador). |

Formicivora consobrina Sclater, P. Z. 8. 1860, p. 279 (1860.—
Babahoyo, S.W. Ecuador); Sclater & Salvin, P. Z. 5. 1879, p. 525
(Pocune).

F, quixensis (nee Cornalia) Cassin, Proc. Acad. N. Sci. Philad.
1860, p. 190 (R. Truando).

No. 1963. gd ad. San Joaquim, Bahia del Chocé, 3.viii.08.—
Wing 48; bill 12 mm.

No. 2142. S$ ad. Sipi: 1.x.08.—Wing 48; tail 45; bill
13 mm.

* Carriker (Ann. Carnegie Mus. vi. nos 2-4, 1910, p. 609), following Mr. Ridgway’s
lead, calls the Costa-Rica form MW. menetriesi schisticolor. As a matter of fact,
however, the bird long known under the name MW. menetriesii has nothing to do with
menetriesii of D’Orbigny, as I have clearly demonstrated in the paper quoted above.
M. menetriesii, D’Orb. belongs to quite a different section, being a close ally of
M. cinereiventris Scl. & Salv. Cfr. Bull. Soc. Philom. Paris (9) viii. 1906, pp. 51-2;
Nov. Zool. xiv. 1907, pp. 69-70.

+ M. sancte-marte Allen, Bull. Amer. Mus. xiii. p. 160 (1900.—Valparaiso,
Santa Marta).

t M. menetriesi (errore) Sclater, Cat. B. xv. p. 240 (part.; 5!, c’).

1164 MR. C. E. HELLMAYR ON THE

Nos. 2277, 2327. ¢@ gad. Novita: 13, 23.x1.08.— Wing 48,
49; tail 47, 46; bill 132 mm.

Nos. 2276, 2310. 9 Q ad. Névita: 13, 19.x1.08—Wing 48;
tail 45; bill 13 mm.

No. 2143. @ juv. Sipi: 1.x.08—Wing 47; tail 47; bill
12 mm.

‘“‘ Tris dark brown, feet and bill black.”

The series agrees with specimens from Western Ecuador. One
of the females (no. 2143) approaches /. g. bowcardi Scl., from
Central America, in the paleness of the under parts, yet it can
easily be distinguished by its much smaller, narrower bill, and
much longer white tips to the outer rectrices.

F. q. consobrina is peculiar to W. Ecuador and W. Colombia,
where it inhabits the hot, forest-covered low country from sea-
level up to about 1500 feet.

122. RAMPHOCENUS CINEREIVENTRIS CINEREIVENTRIS Sel.

Rhamphocenus cinereiveniris Sclater, P. Z.S. 1855, p. 76,
pl. Ixxxvi. (June 1855.—‘ Pasto,” 8. Colombia, Delattre coll.).

No. 2506. ¢ ad. Novita: 29.1.09—Wing 53; tail 31;
bul 18 mm.

No. 2102. 9 ad. Sipi: 21.1x.08— Wing 52; tail 32; bill
19 mm.

No. 1976. ¢g imm. Guineo, Rio Calima: 7.viii.08.—Wing 54;
tail 35; bill 18 mm.

“Tris dark brown, feet blue, maxilla black, mandible grey.” __

In addition we have an adult male obtained by Mr.Hugéne
André at El Paillon, near Buenaventura, May 6, 1899.

The specimens from Western Colombia and fourteen others
from various localities in Western Ecuador (Chimbo, 8. Javier,
Cachyjacu, Lita, Rio Verde) agree perfectly with Sclater’s
original description, having a very distinct, dusky brown post-
ocular streak, and the under parts of a rather clear cinereous
with whitish admixture along the middle line. The type-speci-
men is said to have been obtained at Pasto, 8. Colombia, alt.
8000 feet, which is certainly erroneous. JW. c. cinereiventris 1s
exclusively restricted to the low country and slopes of the
Western Cordillera between sea-level and about 3000 feet.

RR. c. semitorquatus Lawr,*, of which | have examined a good
series from Eastern Costa Rica (Carrillo) and Chiriqui (Boquete),
may easily be distinguished in lacking the brown postocular
stripe and in having the belly rather darker cinereous with very
little, if any, whitish suffusion in the middle,

A third geographical race of this group inhabits Hastern
Ecuadory. Jt resembles the Central American form in the
absence of the dusky postocular streak, but the upper parts are
darker and the sides of the head much deeper coloured. I do not,

* Rhamphocenus semitorquatus Lawrence, Ann. Lyc. N. H. N.Y. vii. p, 469
(1862.—Panama).

+ R. cinereiventris (nec Sclater, 1855) Sclater, Cat. B. Brit. Mus, xv. p, 262
(Sarayacu, East. Ecuador).

BIRDS OF WESTERN COLOMBIA. 1165

however, feel justified in bestowing a name upon this subspecies,
having seen but a few skins in not very good condition.

123. CERCOMACRA TYRANNINA RUFIVENTRIS Lawr.

| Pyriglena tyrannina Sclater, P. Z.S. 1855, p. 90, pl. xeviil.
(1855.—Bogota coll.). |

Disythanunus (sic) rufiventris Lawrence, Ann. Lyc. N. H. N.Y.
vill. p. 131 (1865.—New Granada, line of Panama Railroad—
descr. g juv.); cfr. Salvin, Ibis, 1874, p. 316.

Cercomacra crepera Bangs, Auk, xviii. p. 365 (1901.—Divala,
Chiriqui).

No. 1997.. g ad. Boca de Calima, Rio San Juan, 18.viii1.08.—
Wing 67; tail 58; bill 163 mm.

No. 1981. 9 imm. Guineo, R. Calima: 8.vii1.08.— Wing 60 ;
bill 16 mm.

No. 2387. 9 ad. Noévita: 10.xi1.08.—Wing 59; bill 163 mm.

No. 2742. 9 ad. Pueblo Rico, 5200 ft., 8.1x.08.—Wing 62;
tail 67 (!); bill 165 mm.

“Tris dark brown, feet blue-grey, bill black, mandible horn-
coloured in females.”

These birds strictly belong to the dark western form separated by
Mr. Bangs as C. crepera. Mr. Carriker * has already alluded to
the variability of its characters, basing his conclusions upon Costa
Rica skins. On comparing the series in the Munich Museum,
it becomes at once evident that two fairly marked races can be
distinguished. Adult males from Guatemala, Costa Rica, Chiriqui,
W. Colombia and N.W. Ecuador are, notwithstanding some
individual variation, much darker, more slaty blackish, both on
upper and lower parts than specimens from Bogota (topotypes),
the upper Orinoco, British Guiana, and North Brazil (Para, Rio
Negro). The females also are somewhat darker ferruginous
underneath. The single male from Calima is slightly paler
grey on the belly, but similar specimens I have seen from
S.W. Costa Rica, while several from N.W. Ecuador (San Javier)
in intensity of coloration are practically identical with topo-
typical Chiriqui-skins. Mr. Bangs (J. c. p. 366) refers the birds
from Loma del Leon (= Lion Hill), Panama, to typical tyrannina.
I doubt, however, the correctness of this view, since two males
from Monte Oscuro, near Panama City, though not extreme
examples of crepera, are certainly much nearer that form than to
tyrannina. ‘Therefore, it seems to me that Lawrence’s term
rufiventris, founded upon an immature male from Lion Hill, must
take precedence over crepera.

According to my views, the range of the two subspecies would
be as follows :—

(a) C. tyrannina tyrannina (east of the Andes): E. Colombia
(Bogota, Bucaramanga, etc.); Wenezuela: Orinoco (Munduapo)
and Caura Valley; British Guiana; Northern Brazil: Rio Negro
down to Manaos; Obidos, north bank of Amazons; Para district.

* Ann, Carnegie Mus. v, 1908, p. 8; 7.c. vi. 2-4, 1910, p. 618.

1166 MR. C. E. HELLMAYR ON THE

(b) C. tyrannina rufiventris Lawr. Central America from
Guatemala southwards, Western Colombia and Western Kcuador,
south to Chimbo.

124, CercoMACRA NIGRICANS NScl.

Cercomacra nigricans Sclater, P. Z. 8. 1858, p. 245 (Nov. 1858.—
Santa Marta * (type) and Bogots-coll., Colombia ; descr. ¢ imm.) ;
Sclater & Salvin, P. Z. 8. 1879, p. 526 (Remedios, Antioquia) ;
Berlepsch, Journ. f. Orn. 1884, p. 308 (Bucaramanga).

Pyriglena naculicaudis Sclater, op. at. p. 247 (Nov. 1858.—
“Trinidad,” errore; descr. ¢ ad.).

No. 1989. ¢S ad. Mouth of Calima, Rio San Juan, 13.viii.08.—
Wing 67; tail 67; bill 18 mm.

“ Tris dark brown, feet blue, bill black.”

This bird is in the “ maculicaudis” plumage, viz. glossy black
all over, with the campterium, a large dorsal patch, distinct edges
to the wing-coverts, and long apical spots to the outer rectrices
white, and agrees with other adult males from N.W. Ecuador,
and several Bogota skins. Berlepsch & Hartert have shown
that C. maculicaudis and C. nigricans are merely different sexes
of the same species, and the examination of a large series of
sexed specimens has convinced me that this view is perfectly
right. It should also be borne in mind that the supposed two
species have exactly the same range, extending from the Isthmus
of Panama south to Western Ecuador. Mr. Cherrie took three
specimens at Altagracia, Orinoco, two of which, marked “ g ” by
the collector, are in the ‘“ maculicaudis” phase, while the third,
sexed as “92,” is even greyer, both above and below, than the
type of nigricans. . The locality “Trinidad” given for P. maculi-
caudis is certainly erroneous. I have examined the type and
found it practically identical with adult males from Panama and
Bogota. The preparation furnishes no clue as to its origin. The
specimen appears to have been remade, though it looks somewhat
like the skins imported from Northern Colombia (Baranquilla,
Cartagena, etc.).

C. carbonaria Scl. & Salv.t, from the Upper Rio Branco in
Northern Brazil, is quite distinct, although nearly allied. The
male differs in having the upper parts sooty grey ; the throat and
breast much duller black, shading into sooty grey on the flanks ;
in the shorter white tips of the rectrices, as well as in having
distinct, though narrow, white apical edges to the secondaries.
The female, too, is quite different, having the throat white freckled
with dusky, and the belly bright ochraceous with the middle line
whitish. C.carbonaria also has a much narrower and some-
what shorter bill. So far it has been met with only by Natterer,

* Although not taken in recent years by any collector, C. nigricans may yet be
found in the Santa Marta district, for there is a specimen from Calamar, lower
Magdalena (August 5, 1898) in the collection of H.R.H. The Princess Therese of
Bavaria.

+ Nomencl. Av. Neotrop. p. 161 (1873.—Rio Branco, N. Brazil).

BIRDS OF WESTERN COLOMBIA. 1167

who obtained a large series at Forte do Joaquim, on the confines
of British Guiana.

125. CERCOMACRA BERLEPScHI Hart.

Pyriglena berlepschi Hartert, Bull. B. O. C. vii. p. xxix (1898.—
Cachabi, N.W. Ecuador; = ¢ ad.).

Thamnophilus cachabiensis Hartert, 1. c. p. xxix (1898.—
Cachabi, N.W. Ecuador; = @).

Cercomacra berlepschi Hartert, Nov. Zool. ix. 1902, p. 612
(crit.).

No. 1958. g ad. 8. Joaquim, Bahia del Chocd, 1.vi1.08.—
Wing 68: tail 46; bill 194 mm.

No. 2089. gad. Near Sipi, 200 feet alt., 18.1x.08.— Wing 69 ; |
tail 474; bill 19 mm.

No. 1966. Q@ ad. 8S. Joaquim, 4.vii.08.— Wing 67; tail 43 ;
bill 19 mm.

‘‘ Tris dark red, feet and bill black.”

The males are uniform deep black, with a large, concealed
white interscapular patch. The female is rather duller black,
the abdomen slate-blackish, and the feathers of the throat, fore-
neck, middle of the breast, as well as the lesser and median upper
wing-coverts, show very distinct, though sometimes slight, white
apical spots or edges. Dr. Hartert has already pointed out that
these two types of coloration, originally described as different
species, were merely male and female of one and the same species.
From the evidence at hand, there can be no doubt that this view is
correct.

The specimens from Chocé agree in every respect with the
series from N.W. Ecuador in the Tring Museum*. (C. berlepschi
is somewhat difficult to place. In general form it closely re-
sembles some of the short-tailed species, e. g. C. tyrannina,
but the tail is much shorter and slightly rounded instead of
being graduated, the wing relatively longer, and the bill longer
as well as heavier. Although it might some day be found
necessary to create a new genus for its reception, it would be
unwise to do so without a thorough study of all the related
groups.

126. HypocnemMis N&ZVIOIDES Lafr.

Conopophaga nevioides Lafresnaye, Rev. Zool. x. p. 69 (1847—
no locality given; type in Mus. Acad. Philadelphia, coll.
Delattre—we fix as type locality Panama whence the Derby
Museum (Liverpool) possesses two specimens obtained by De-
lattre ; cfr. Sclater, P. Z.S. 1858, p. 254).

Hypocnemis nevioides Cassin, Proc. Acad. N. Sci. Philad. 1860,
p. 190 (Falls of the Truando).

* In this connection a misleading error may be corrected. Hartert (7. c.) gives
the length of the wing for P. berlepschi, as 44 to 46 mm. However, five adult males
from N.W. Ecuador, including the type, in the Tring Museum present the following
dimensions ; wing 67-683 ; tail 48-47; bill 19-20 mm. The females (5) are slightly
smaller; wing 64-68 ; tail 41-46; bill 18-19 mm.

1168 MR. GC. E. HELLMAYR ON THE

No. 2603. g ad. Condoto: 16.iv.09.—Wing 638; tail 38;
bill 16 mm. ;

Nos, 2341, 2491. 9 Q ad. Novita: 26.x1.08, 25.1.09.— Wing
62, 61; tail 37, 34; bill 16 mm.

Nos. 1977, 1999. 2 9 imm. Guineo: 7.viii; mouth of Caiima ;
18.vi1i.08.— Wing 64, 62; tail 37, 35; bill 16 min:

“Tris brown, feet blue-grey, bill Tage, lower mandible of
females grey.”

These specimens as well as others from N.W. Eeuador (San
Javier) agree perfectly with typical Panama examples. Skins from
Miravelles, Costa Rica, have, as a rule, the wing a trifle larger,
and have the back of a clearer, brighter chestnut, but these
variations are so iInconstant that it appears to me highly im-
probable that the northern form, H. n. capnitis Bangs* can be
maintained. This view is shared by Mr. Carriker 7, who collected
a large series of this species in Costa Rica.

Birds from different localities measure as follows :—

Five adult males from Miravelles, Costa Rica: wing 66-67;
tail 34-37 mm.
Two adult males from Panama: wing 65, 653; tail 36, 37 mm.
One adult male from W. Colombia: wing 63; tail 38 mm.
Three adult males from N.W. Ecuador (8. Javier): wing
63-65; tail 35-374 mm.

Four adult females from Miravelles, Costa Rica: wing
63-65 ; tail 33-36 mm.

Two adult females from W. Colombia: wing 62, 61; tail 37,
o4 mm.

One adult female from N.W. Ecuador (S. Javier): wing 61 ;
tail 36 mm.

Both Mr. Bangs and Mr. Carriker give ‘“‘ Pasto, Cauca valley,
Colombia” as the type locahty, but this is an obvious error.
First of all, Lafresnaye does not say where his type came from,
and secondly the occurrence of H. nevioides in the vicinity
of Pasto (which hes high up in the mountains at an elevation
of nearly 8000 feet) is utterly impossible, the species being
restricted to the humid, tropical lowlands, from sea-level to about
2000 feet. Delattre most probably obtained the type at
Panama, whence there are two of his skins in the Derby Museum
(see above).

127. MyYRMELASTES IMMACULATUS IMMACULATUS Lafyr.

Thamnophilus immaculatus Lafresnaye, Rev. Zool. viii. p. 340
(1845—“‘ad Bogotam,” descr. ¢ 2).
_ Myrmeciza berlepschi (nec Ridgway) Bangs, Proc. Biol. Soc.
Wash. xxiii. p. 73 (Palmar, Pavas, La Maria, W. Colombia).
No. 2818. g ad. Pueblo Rico, San Juan slopes, 28.x.09.—
Wing 84; tail 77; bill 22 mm.
* Proc. Biol. Soc. Wash. xix. p. 107 (1906.—Miravelles, Costa Rica).
7 Aun. Carnegie Mus. vi. 1910, p. 619.

=

BIRDS OF WESTERN COLOMBIA. 1169

[Tring Museum. @ ad. . Primavera, Cordillera occidental,

1904. Raap coll. No. 390.—Wing 79; tail 79; bill 20 mm. }

Mr. Ridgway * has correctly pointed out the distinctness of
M. immaculatus, from Bogota, and IM. 2. berlepschi Ridgw., from
Western Ecuador, though some of his conclusions prove to be
not well-founded. First of all, d/. 2. berlepschi does not occur
anywhere in Colombia, the specimens from Bogota mentioned by
Berlepsch & Taczanowski t, which were kindly forwarded to me
by the Count, being clearly referable to true IZ. immaculatus. The
female from Primavera, W. Colombia, is practically identical with
several Bogota skins in the Berlepsch Collection, In all these
females the lower parts (except the blackish upper throat) are
rufescent brown, mottled with dull smoky grey on the foreneck
and in the middle of the breast, while there is only a narrow
white stripe on the bend of the wing.

Sixteen females from Western Ecuador (Chimbo, Paramba,
Lita, Gualea, Bulan, ete.) have a much larger, heavier bill, the
forehead is more scantily feathered, the bend of the wing shows
a large white patch, and the lower parts are much brighter
rufous brown. The chin as well as the sides of the head are
blackish, exactly as in J/. immaculatus from Colombia. Seventeen
adult males from Western Kcuador also differ from the Colombian
ones in their larger bill, more scantily feathered forehead, and
in having much more white on the shoulders. The general
plumage is sometimes, though not always, deeper black.

M. immaculatus zeledoni Ridgw. t, lately united to the Colom-
bian form by Mr. Carriker§, appears to me to be much more
closely allied to IM. 7. berlepschi. In fact, on comparing three
adult males and two females from Costa Rica (Cariblanco de
Sarapiqui) with the large series from Keuador, I find the differences
not very pronounced. All that can be said is that the northern
birds have the white shoulder-patch slightly smaller. The females
otherwise agree with those from Keuador. More material from
Costa Rica may even show the two forms to be inseparable.

128. MyYRMELASTES EXSUL MACULIFER Hellm.

[Myrmeciza exsul Sclater, P. Z. 8. 1858, p.540 (1859.—Panama:
Delattre). }

Myrmelastes exsul maculifer Hellmayr, Nov. Zool. xiii. p. 340
(1906.—Paramba, N. Ecuador).

Myrmeciza exsul (nec Sclater) Cassin, Proc. Acad. N. Sci. Philad.
1860, p. 191, no. 98 (= 3), 99 (= 2) (Turbo); Sclater & Salvin,
P.Z.8. 1879, p. 526 (Rio Neche, Antioquia).

Myrmelastes cassini Ridgway, Proc. Biol. Soc. Wash. xxi. p.194
(1908.—Turbo, N. Colombia).

* Proc. Biol. Soc. Wash. xxii. 1909, p. 74.

+ P. Z. S. 1883, p. 565.

£ Myrmeciza zeledoni Ridgway, Proc. Biol. Soc. Wash. xxii. p. 74 (1909.—
Guayabo, E. Costa Rica).

§ Ann. Carnegie Mus. vi. 1910, pp. 618-9.

1170 MR. C. E. HELLMAYR ON THE

Nos. 2082, 2145, 2178. g$ dad. Sipi: 12.ix, 2, 12.x.08.—
Wing 67-69; tail 444-47; bill 18 mm.

No. 2234. gad. Rio Cajén: 3.xi.08.—Wing 68; tail 45;
bill 18 mm.

No. 2488. gad. Névita: 28.x11.08.—Wing 69; tail 433 ; bill
19 mm.

Nos. 2017, 2018, 2038. 9 2. Noanama: 26, 31.viii.08.—
Wing 63; tail 40-423; bill 173-18 mm.

Nos. 2179, 2353) 9) adi, od) juv.. )\Sipiy 12x08); INovatare
30.xi.08.— Wing 63, 66; tail 42, 44; bill 18, 19 mm.

“Tris dark red, feet grey, bill black.”

When discussing the status of this species and its northern
limits I alluded to the paler. less rufescent brown coloration of the
specimens from Neche, Northern Colombia, in the British Museum
(cfr. Nov. Zool. xiii. p. 342), and two years later Mr. Ridgway
separated two similar skins from Turbo under the name of
M, cassini*. The present series, however, shows considerable
variation in the colour of the back. Six of the specimens have
the back of exactly the same deep rufous or vandyke- brown shade
as a large series from N.W. Ecuador, while the four remaining
ones are somewhat lighter, more mars-brown above, like the two
examples from Neche, Antioquia. Two adult males and one
female obtained by Mr. Rosenberg on the Rio Dagua in 1895,
agree again with the types from Paramba.

Neither is there any constant difference in the coloration of the
head and underparts. Four males—two from Paramba, one from
Sipi, one from Névita—are decidedly slaty blackish, while the
majority of the skins, both from W. Ecuador and from Choco,
have the chest and belly paler, slate-grey.

Therefore, I cannot admit d/. cassini as a valid form unless
the Turbo birds be distinguished by other characters than those
given in the original description. In addition to the above ten
specimens, the Munich Museum possesses seven males and four
females from N.W. Ecuador; and I have examined ten more
from W. Ecuador, three from the Rio Dagua, in the Tring
Museum, and the couple from Neche in London.

M. e. maculifer, as understoood by me, is another Pacific
type, its range being limited to the forest districts of Western
Ecuador and Colombia.

129. ANOPLOPS BICOLOR DAGU& Hellm.

[Pithys bicolor Lawrence, Ann. Lyc. N. H.N. Y. vii. p. 6
(May 1863.—Panama Railway). |

Gymnopithys bicolor dague Hellmayr, Bull. B.O. C. xvi. p. 83
{1906.—EI Paillon near Buenaventura, Chocé, W. Colombia ; type
in Tring Mus.).

* Mr. Ridgway also mentions a female from Cascajal, Coclé (Panama), as
belonging to this form. There must, however, be some mistake with regard to the
locality, since three adult males from that place in the Tring Museum are clearly
referable to M. easul exsul. Heyde’s localities are not always trustworthy, and, as

he also got many specimens from Western Colombia (Ndévita etc.), an error in
labelling the bird might have occurred.

BIRDS OF WESTERN COLOMBIA. 1171

No. 2336. gad. Noévita: 25.xi.08.—Wing 78; tail 51; bill
20 mm.

Nos. 2564, 2565. ¢ 2 ad. Juntas, R. Tamand: 9.111.09.—
Wing 77, 76; tail 48, 49; bill 20, 193 mm.

“Tris brown, feet blue-grey, maxilla black, mandible grey.”

The specimens are topotypical. The female differs from the
males in having the dark grey colour of the forehead and sides of
occiput less extended and in lacking the blackish border to the
white chest.

A. b. dague is nearest to A. b. bicolor, from Panama, but may
be easily distinguished by the much darker, deep vandyke (rufous)
brown instead of light russet-brown colour of the upper parts,
much darker rufous-brown flanks, larger, stronger bill, and by
the presence of a distinct blackish border to the white chest in
the male sex.

A. b. equatorialis Hellm., from Western Ecuador, agrees with
A. b. daguwe in the dark coloration of the back and flanks, but the
forehead and crown are bright cinnamon rufous, the sides of the
occiput alone being dark cinereous, etc.

The various races of A. bicolor not having been properly
understood hitherto, it is hoped that the subjoined key will aid
ornithologists in identifying specimens of the group. A. leucaspis
Scl. is a near ally of A. bicolor and very likely its Amazonian
(eastern ) representative ; it differs, however, in having the cheeks,
malar region, and pntecion ear-coverts white (instead of black), and
in the possession of a large, concealed, pale cinnamomeous inter-
scapular patch in the female sex.

Key to the species and subspecies.

1, Cheeks, malar region, and anterior ear-coverts white.
Female with a distinct, light cinnamomeous inter-
scapular blotch. {Nasal plumes and whole pileum
bright ferruginous. Lores and narrow sabeenaty
stripe black]... A. leucaspis.

— Cheeks. malar region, “and whole of the ear-coverts
black. Female ‘without a pale interscapular blotch 2.

2. Sides of the occiput dark cinereous, Av Sie con-
trasted with colour of crown ...... 3.

— Sides of the occiput concolor with crown, never
cinereous. [White chest laterally not bordered aby

a blackish stripe]... 5.
3. Forehead and a broad ‘superciliary stripe dark cine-

reous like the sides of the occiput ...... 4.
— Forehead and crown ferruginous, only sides of ‘occiput

dark cinereous ...... : A. bicolor equatorialis.
4. Crown and back light yusset- brown ; ‘sides of body

mars-brown; bill “smaller : 17-183 mm. 58 A. bicolor bicolor.

— Crown and back deep vandyke or dea rufous brow n;

sides of body vandyke brown; bill larger;

ICO arab eee boSenR a eRetpeocuede sameogoncs saecaeend manceds A. bicolor dague.
5. Head above light russet-brown; back umber-brown

or very slightly tinged with rufescent; sides of

body light mars-brown .............0....eee cee cee cece eee A. bicolor olivascens.
— Head above bright ferruginous ; back decidedly cinna-

mon-brown; sides of body rufescent or russet-brown 4. bicolor ruficeps.

1172 MR. C. E. HELLMAYR ON THE

(1) ANopLops LEUCASPIS Scl.

Myrmeciza leucaspis Sclater, P. Z. 8. 1854, p. 253, pl. Ixx.
(1855.—Bogoté, Colombia (type); Chamicuros, Hast. Peru ;
Cobati, Rio Negro).

Pithys leucaspis Sclater & Salvin, P. Z. 8. 1867, p. 576 (Cobati) ;
lidem, I. c. p. 751; iidem, |. c. 1873, p. 276 (Xeberos, Chyavetas,
E. Peru) ; Pelzeln, Zur Orn. Bras. ii. p. 89 (Barcellos, R. Iganna,
R. Uaupés: R. Negro); Taczanowski, Orn. Pér. ii. p. 74 (Tarapoto,
N. Peru).

Gymnopithys leucaspis Salvin & Godman, Biologia, Aves, 11.
p. 222 (Rio Meta, Hast. Colombia).

This species is exclusively found in the forests of Upper
Amazonia, from the eastern slopes of the Colombian Andes
south to Northern Peru. Both Natterer and Wallace obtained it
on the banks of the Rio Negro, while Wheeler transmitted specimens
from the Rio Meta, E. Colombia, to the British Museum. Haux-
well met with it at Chamicuros, Eastern Peru, and KE. Bartlett
procured a series at Chyavetas, Xeberos, and Chamicuros, now in
the Tring Museum. A. leucaspis is also occasionally found in
the trade-collections sent from Bogota to Europe.

Against the statement in the ‘ Cat. of Birds,’ xv. p. 295, I wish
once more to emphasize that it is the female which possesses the
cinnamomeous interscapular blotch. Five sexed males from
Northern Peru (Chamicuros, Chyavetas, Xeberos ; Bartlett coll.)
in Mr. Rothschild’s collection, five from the Rio Negro, obtained
by Natterer, in the Vienna Museum, and one male from Cobati,
R. Negro, Wallace coll. in the British Museum, show no trace of
it; while in five sexed females from the Rio Negro (Natterer ;
Vienna Museum) there is a large conspicuous patch at the base
of the dorsal feathers. Besides the above, I have examined two
males and four females from “ Bogota-coll.” in the collections at
Tring, Munich, and Paris.

(2) ANOPLOPS BICOLOR #QUATORIALIS Hellm.

Pithys bicolor equatorialis Hellmayr, Orn. Monatsber. x. p. 33
(1902.—Lita, N.W. Ecuador).

P. leucaspis (nec Sclater) Hartert, Nov. Zool. v. 1898, p. 495
(Chimbo, W. Ecuador); Goodfellow, Ibis, 1902, p. 65 (Sto.
Domingo, W. Ecuador).

Gymnopithys ruficeps (nec Salvin & Godman) Salvadori & Festa,
Boll. Mus. Zool. Torino, xiv. no. 362, 1899, p. 32 (Santiago,
W. Ecuador); Hartert, Nov. Zool. ix. 1902, p. 613 (Chimbo,
Paramba, 8. Javier, W. Eeuador).

This form, which is confined to Western Ecuador, shares with
A. leucaspis the bright ferruginous colour of the forehead and
crown, but is much darker rufous brown on the back, has the
sides of the occiput dark cinereous, etc., ete. I have examined
twenty specimens from W. Ecuador in the Tring, Vienna, and
Munich Museums. There is never any trace of a pale inter-
scapular patch.

BIRDS OF WESTERN COLOMBIA. 1a

(3) ANOPLOPS BICOLOR RUFICEPS Salv. & Godm.

Gymnopithys ruficeps Salvin & Godman, Biologia, Aves, ii. p. 222
(1892.—* Cauca Valley,” the types are from Neche and Remedios,
Antioquia).

Pithys leucaspis (nec Sclater) Sclater & Salvin, P. Z. 8. 1879,
p- 526 (Remedios, Neche).

This race is evidently confined to the Cauca Valley in the
province of Antioquia, N. Colombia. Besides three of Salmon’s
skins in the British Museum (2) and at Tring (1), I have examined
five Bogota skins * referable to the same form. A. 0. ruficeps
differs at a glance from the preceding subspecies in lacking the
cinereous colour on the sides of the occiput and in the white of
the chest being laterally not bordered with black. It is much
more closely related to A. b. olivascens, but of a more rufescent
colour throughout.

(4) ANopLops BICoLoR DAGUz Hellm.

This well-marked form also has a very limited range, being
known only from the valleys of the Dagua and San Juan Rivers
in Western Colombia. The Tring Museum possesses an adult
male obtained by E. André near Buenaventura, and another from
Juntas, Raap coll. Palmer’s three specimens are the only others
on record. For characters see above.

(5) ANOPLOPS BICOLOR BICOLOR Lawr.

Pithys bicolor Lawrence, Ann. Lye. N. H. N.Y. viii. p. 6 oe
—Panama Railway).

The “typical” subspecies also has a rather limited distribution,
being restricted to Panama and Veragua. I have examined four
skins from the Panama Railroad (McLeannan), one from Chepo,
and one from Santa Fé, Veragua (Arcé), all in the British
Museum.

(6) ANOPLOPS BICOLOR OLIVASCENS Ridgw.

Pithys bicolor olivascens Ridgway, Proc. U. S. Nat. Mus. xiv.
1891, p. 469 (1892.—Santa Ana, Honduras).

This form is more widely distributed, ranging from Hastern
Honduras through Nicaragua, and Costa Rica to Chiri iqui. There
are large series from various localities in the Tring and Munich
Museums.

130. FoRMICARIUS ANALIS DESTRUCTUS Hart.

[Myothera analis Lafresnaye & D’Orbigny, Syn. Av. 1., in Mag.
Zool. el. ii. p. 14 (1837.— Yuracarés, N.E. Bolivia). |

Formicarius analis destructus Hartert, Nov. Zool. v. p. 493
(1898.—Paramba, N.W. Ecuador); idem, 1. c. ix. 1902, p. 614
(N.W. Ecuador; crit.).

F. analis (nec Lafr. & D’Orb.) Sclater, P.Z.S. 1860, p. 294

* One in Tring, two in the Paris Museum, two in the British Museum.

Proc. Zoot. Soc.—1911, No. LX XIX. T®

1174 MR. C; E. HELLMAYR ON THE

(Esmeraldas, N.W. Ecuador); idem, Cat. B. Brit. Mus. XV. p- oe
(part. : Balzar, Santa Rita ; Esmeraldas: W. Keuador).

I. nigr icaypillus (nec Ridgway) Ridgway, Proc. U.S. Mus. xvi.
p- 675 (part.: sp. ew Santa Rita).

F.. destructus Salvadori & Festa, Boll. Mus. Zool. Torino, xiv.
no. 362, 1899, p. 32 (Peripa, W. Ecuador).

Nos. 2508, 2393. ¢ g ad. Névita: 11.xi1.08, 30.1.09.— Wing
90, 86; tail 55, 52; bill 20, 21 mm.

No. 2372. 9 juv. Névita: 5.xi1.08.—Wing 86; bill 18 mm.

‘Tris dark brown, feet and bill black.”

Perfectly identical with topotypical specimens from Paramba,
N.W. Ecuador, the head all round and the hind-neck being sooty
black, abruptly contrasting with the dark rufous brown of the
back.

F. a, destructus is most nearly allied to /. a. nigricapillus Ridgw.
of Eastern Costa Rica. Dr. Hartert * having clearly pointed out
the differences between the two races, I need not dwell any more
upon the subject.

This Yormicarius is confined to the lowlands and foot-hills of
the Pacific coast of Colombia and Ecuador. On the other side of
the Western Cordillera, in the Cauca Valley, another species
or subspecies, /. satwratus Ridgw. tT, takes its place. I have
examined the example from Remedios (Salmon) in the British
Museum, and an adult male obtained by the late J. H. Batty
at Los Jambos (Cauca Valley) in Mr. Rothschild’s Museum at
Tring, which I cannot distinguish from typical Trinidad specimens.
F. satwratus is much paler grey underneath, has the sides of the
neck distinctly cinnamomeous, and the top of the head light
olivaceous brown like the back, besides some minor differences.

131. FoRMICARIUS RUFIPECTUS RUFIPECTUS Salv.

fF. rufipectus Salvin, P. Z. 8. 1866, 73, pl. viii. (1866.—
Santiago de Veragua) ; Salvadori & ines, Boll. Mus. Zool.
Torino, xiv. no. 362 1899, p. 33 (Gualea, W. Keuador) ;
Ménégaux & Hellmayr, Bull. Soe.) Philom: Paris, (9) viii. 1906,
p- 52 (between Ksmeraldas and Pachijal, Oyacachi, W. Keuador) ;
Carriker, Ann. Carnegie Mus. vi. 1910, p. 626 (Rio Cali, W.
Colombia).

No. —. d ad. Pueblo Rico: 16.viii.09.—Wing 99; tail 63;
bill 20 mm. :

No. 2729. 9 ad. Pueblo Rico: 16.viii.09.—Wing 94; tail 59;
bill 214 mm.

“Tris and feet dark brown, bill black.”

These specimens agree well with others from Chiriqui (A. Lara
coll.). The female differs from the male, in addition to its
smaller size, by having the breast somewhat clearer rufous, and

* Nov. Zool. ix. p. 614.

+ Proc. U.S. Mus. xvi. p. 677 (1894.—Trinidad (type) ; Venezuela: S. Esteban ;
N.E. Colombia: Remedios).

BIRDS OF WESTERN COLOMBIA. 1175

the crown dull brownish black, the hind neck only bemg washed
with chestnut, while in the male all the top of the head, from
the forehead to the nape, is dark chestnut-maroon. The sexual
differences noticed by Mr. Carriker, however, do not hold good.
Both specimens have the middle of the belly extensively ochraceous
buff and the flanks dark smoky grey.

This scarce species ranges from Central Costa Rica (Juan
Vinas) to Western Ecuador *, On the eastern (Amazonian)
slopes of the Ecuadorian Andes and in Peru, /. r. rufipectus is
replaced by /. r. thoracicus Tacz. & Berl. f, at once recognizable
by its deep black pileum. I have examined one of the typical
specimens and an adult male from Huaynapata, both in Count
Berlepsch’s collection.

132. PrrrAsoMA ROSENBERGI Hellm.

P. rosenbergi Hellmayr, Rev. Frane¢. d’Orn. ii. p. 51 (April
1911.—Sipi, Rio Sipi).

3 ad. Pileum and nape deep cinnamon-rufous;_ back and
scapulars dull olive, more brownish on the rump, each feather
broadly bordered with black on either side, giving the upper parts
a striped appearance; some of the interscapular feathers white at
the base; upper wing-coverts sepia-brown, washed with russet,
especially on the outer webs, each feather with a small, but
distinct, whitish apical spot, and with a distinct black margin
round the tip; primary coverts uniform blackish brown; remiges
blackish, with the outer web dull russet-brown ; rectrices dusky,
the central ones washed with russet. Lores and broad super-
ciliary stripe, reaching to the sides of the neck, uniform deep
black ; cheeks, subocular and malar region, ear-coverts, chin and
throat ochraceous, rather deeper on the sides of the head; fore-
neck and sides of the breast and flanks dull olive-brownish ;
middle of the breast and abdomen uniform buffy white, this
colour being separated from the ochraceous throat by the dull
brown crescent of the foreneck; under tail-coverts hair-brown,
edged with whitish; axillaries and under wing-coverts dusky
brown, the under primary-coverts tipped with white.

“ Tris dark brown, feet and bill black.”

Wing 97; tail 34; tarsus 45; bill 242 mm.

Type in the Zoological Museum, Munich, No. 09.5759. ¢ ad.
Sipi, Rio Sipi, alt, 150 feet, Sept. 25, 1908. Mervyn G. Palmer
coll. no. 2120.

This species, which is named in honour of Mr. W. F. H.
Rosenberg, of London, who organized Mr. Palmer’s expedition to
Western Colombia, is unfortunately represented by a single spe-
cimen only. ‘The type is a perfectly adult male, as shown by the

* The specimen in the British Museum said to be from Baeza (Eastern Ecuador)
either belongs to F’. x. thoracicus, or else the locality is wrong (Buckley !).

+ F. thoracicus Taczanowski & Berlepsch, P. Z. S, 1885, p. 101 (Machay,
Eastern Ecuador); Salvadori & Festa, Boll. Mus. Zool. Torino, xiv. no. 362, p.33 (S,

José, E. Kcuador); Berlepsch & Stolzmann, Ornis, xiii. 2. 1906, p. 118 (Huavnapata,
S.E. Peru),
Oke

1176 MR. C, E. HELLMAYR ON THE

uniform blackish primary coverts, the whitish apical spots to the
wing-coverts, the black superciliary stripe, etc. Although closely
allied to P. rufopileatum Hart.*, from North-western Ecuador,
it differs from both sexes of that species ft in having no dusky
eross-bands whatever on the lower parts. In P. rufopileatuwm the
male has the under surface of the body, with the exception of
the flanks, regularly barred with black and white, the black
markings of the throat being often broken and less pronounced,
though always present, while only the sides of the head are
ochraceous. The superciliary stripe (uniform black), back and
upper wing-coverts are exactly asin P. rosenbergi. The female
ot P. rufopileatum agrees with the latter in the ochraceous throat,
but can, of course, be easily distinguished by having the feathers
of the superciliary region white, edged with black, and the lower
parts (posterior to throat) bright ochraceous narrowly banded
across with dusky olive. Besides, the back is much more decidedly
brown, and the apical spots to the wing-coverts are bright: buff
instead of white.

133. CoNOPOPHAGA CASTANEICEPS CASTANEICEPS Scl.

Conopophaga castaneiceps Sclater, P. Z. 8. xxiv. p. 47 (1857—
part., type from Bogota, Colombia; = ¢).

C. gutturalis idem, |. c. 1868, p. 574 (Bogota; = @).

No. 3779. @ ad. La Selva, Rio Jamaraya, 4600 ft., 1.x.09.—
Wing 72; tail 39; bill 143 mm,

‘“‘Tris dark brown, feet grey, maxilla black, mandible grey.”

This bird corresponds with Sclater’s description of C. gutturalis.
Tt has the back rather more brownish than a Bogota skin, but
does not differ otherwise. This is, so far as I know, the first
record of C. castaneiceps from the Western Cordillera of Colombia.

In Peru it is replaced by a closely allied form, C. c. b#uwnneinucha
Berl. & Stolazm.f The female of C. c. castaneiceps, in general
coloration, is much hike Grallaricula cucullata Scl., but may
easily be recognized by its white postocular stripe (‘‘ pencil”).

134, ANDRODON ZQUATORIALIS Gould.

Androdon equatorialis Gould, Ann. Mag. N. H. (3) xii. p. 247
(1863. — Ecuador) ; Sclater & Salvin, P. Z. 8. 1879, p. 528
(Remedios, Antioquia).

No. 2129. g ad. Sipi, Rio Sipi: 29.ix.08.—Wing 67; tail
42; bill 40 mm.

Nos. 2074, 2644. 6 d imm. Sipi: 10.1x.08; Tadd: 8.v.09.—
Wing 68, 69; tail 42, 44; bill 37, 40 mm.

“Tris black, feet pink, maxilla black, mandible light yellow.”

The first specimen (No, 2129) has the forehead and crown

* Novit. Zool. viii. p. 370 (1901.—Bultin, N.W. Kcuador).

+ The bird figured in Nov. Zool. ix. 1902, pl. viii. fig. 2 as “juv.” is really
the adult female. This is quite evident from the large series, partly im the Tring

Museum, partly in the possession of Mr. Rosenberg, which I have examined.
t P. Z.S. 1896, p. 385 (Garita del Sol, La Gloria, Chanchamayo, C. Peru).

BIRDS OF WESTERN COLOMBIA. ILI

shining coppery red, passing into golden green on the occiput; in
the two other, apparently immature, birds the forehead is dull
greyish, while the feathers of the crown are dusky, tipped with
bronze-green and edged with reddish bronze. No. 2074 shows a
strong bluish tinge on the nape, completely absent in the other
examples,

With regard to the variation of this species, cfr. Simon, Bull.
Mus. Paris, 1907, no. 1, p. 17.

A. equatorialis is strictly confined to the lowlands and hill-
ranges of Western Ecuador and Colombia, ranging from near
sea-level up to about 2400 feet.

135, THRENETES RUCKERI FRASERI Gould.

[Trochilus Ruckert Bourcier, P.Z.8. 1847, p. 46 (May 1847—
loc. ign.) *.]

Giaucis frasert Gould, Monogr. Trochil. pt. xxiv. pl. 12 (1861.
—KHEsmeraldas, N.W. Ecuador).

Threnetes frasert Hartert, Nov. Zool. v. 1898, p. 493 (Cachabi,
N.W. Ecuador) ; Bangs, Proc. Biol. Soc. Wash. xxiii. 1910, p. 72
(Naranjito, Rio Dagua ; Palmer coll.).

Threnetes ruchert (sic) Boucard, ‘ Humming Bird,’ v. 1895, p. 7
(Rio Dagua).

No. 2417. $ fere ad. Noévita, 150 ft., 19.xii.08.— Wing 60;
tail 38; bill 31 mm.

“ Tris black, feet pink, maxilla black, mandible yellow.”

The differences separating 7’. r. ruckert and T. 7. fraseri are so
slight that their relationship is better expressed by trinomials.
One of the principal characters given by authors for the southern
bird, namely, the lesser extent of the cimnamon-rufous patch on the
foreneck, is of no value at all. Adulé males of fraseri have quite
as much, sometimes even more, rufous on the jugulum than
Central American skins. The only constant points by which to
tell frasert are the rather darker smoke-grey belly, the pure
metallic green upper parts, without any bronzy or golden gloss,
and the much deeper, decidedly bluish-black colour of the median
portion of the lateral rectrices.

Specimens from Western Hcuador have the two central
rectrices dull green, passing to nearly blackish green towards the
tip, while in the male from Névita they are much brighter, uniform
metallic green. In that respect the Choco bird slightly points
towards ruckert from Central America, but otherwise it is
perfectly typical of fraseri.

T. r. fraseri inhabits the forest-covered lowlands of Western
Colombia and Western Ecuador‘, from sea-level up to about
700 feet.

* Bourcier’s rather vague description is possibly not referable to the present species.
The words “gorge et dessous du corps gris-noir bronzé ”’ can scarcely be applied to
T. ruckeri auct., and a re-examination of the type in the Loddiges Collection
seems desirable.

+ Its reported occurrence in Eastern Ecuador (Sarayacu: cfr. Cat. B. xvi. p. 266)
rests on two of Buckley’s skins, no doubt incorrectly labelled.

.

1178 MR. C. E. HELLMAYR ON THE

136. GLAUCIS HIRSUTA ZNEA Lawr.

| Trochilus hirsutus Gmelin, Syst. Nat. 1, i. p. 490 (1788—ex
Marcgrave: East Brazil). |

Glaucis eneus Lawrence, Proc. Ac. N. Sci. Philad. xix. p. 232
ee fie Vor Rica).

G. columbiana Boucard, Gen. Hum. Birds, p. 402 (end of 1895.
eau Dagua, W. Colombia) ; ; idem, ‘The Humming Bird,’ v. p. 7
(Rio Dagua).

G. hirsuta (nec Gmelin) Sclater & Salvin, P. Z. 8. 1879, p. 528
(Santa Elena).

No. 1985. ¢ (2) ad. Guineo, Rio Calima: 10.viii.08.— Wing
d4L; tail 34; bill 29mm.

a ie is black, feet yellow, maxilla black, mandible Bllevk
grey.”

I agree with Mons. Simon* that the earliest available name
for the small, dark-coloured form of the Pacific coast is the one
given above. Jam unable to find any constant differences, either
in size or in colour, between examples from N.W. Ecuador and
W. Colombia, on the one hand, or those from Central America on
the other.. The uniform cinnamon-brown colour of the under
parts is by no means a constant character of the Western birds,
as already pointed out by Mr. Oberholser ‘.

G. h. cwnea ranges from Costa Rica and Panama south to
N.W. Hcuador (province of Esmeraldas).

137. PH#THORNIS YARUQUI SANCTI-JOHANNIS Hellm.

| Trochilus Yaruqui Bourcier, Compt. Rend. Ac. Sc. Paris, xxxil.
No. 6 (séance 10 Febr. ), p: 187 (1851.—*‘ Les bois tres-chauds
des environs d’Yaruqui,’ W. Kcuador). |

Phaéthornis yaruqui sancti-johannis Hellmayr, Bull. B. O. C.
xxvii. p. 92 (1911.—Condoto).
~ Phethornis yaruqui (nec Bourcier) Cassin, Proc. Acad. N. Sci.
Philad. 1860, p. 194 (R. Truando); Simon & Dalmas, Ornis, x1.
p. 218 (Buenaventura).

Nos. 2128, 2197, 2344, 2598 +. -$ gd ad., ¢ nearly ad. Sipi:
929.ix.; Noanama: 17.x.; Névita: 27.x1.08.; Condoto: 14.iv.09.
—Wing 61-62; tail 55-61; bill 41-43 mm.

Nos. 1987, 2042, 2095. 92 ad, @ 2 imm. Guineo: 11.viil.;
Noanama: 3l.vili.; near Sipi: 19.1x.08.—Wing 58-59; tail
59-63 ; bill 37-40 mm.

“Tris black, feet brown, maxilla black, mandible scarlet.”

Type of subspecies in Zoological Museum, Munich: no. 09.
5807. dad. Condoto, R. Condoto, 150 feet, April 14, 1909.
M. G. Palmer coll. no. 2598.

P. y. sancti-johannis is a very distinct form, differing in many
* Rey. Frang. d’Orn. i. 1910, p. 260.

- + Proc. U.S. Mus. xxiv. 1902, p. 311.
{ Type of subspecies.

BIRDS OF WESTERN COLOMBIA. 1179

important points from P. y. yaruqui of Western Ecuador. The
males may be distinguished by their shorter bill; much broader
and darker, deep ochreous-buff postocular and malar streaks ; and
by having the pale smoke-grey median stripe of the throat and
foreneck much wider and confluent with the smoke-grey abdomen,
which is but slightly spotted with metallic green on the flanks ;
whereas in P. y. yaruqui the under surface is mainly dark metallic
green, with the abdomen and a short stripe along the middle of
the throat sooty blackish. The Chocé birds have, too, the base
of the outer web of the rectrices decidedly washed with greenish.
The females are even more different from the Keuadorian yaruqut.
The under surface is light mouse-grey, the sides of the foreneck
and breast being but sparingly mottled with metallic green, and
the pale median stripe along the throat, foreneck, and chest is
conspicuously mixed with whitish. The deep ochreous-buff post-
ocular stripe is much broader, the white tip to the central rectrices
more extended, but less sharply defined than in the females of
P.y. yaruqui. In the females of both races, the malar streak 1s
white, anteriorly tinged with buff.

Mons. Simon, of Paris, to whom I have submitted our series,
agrees to the distinctness of the Colombian race.

P. y. sancti-johannis replaces the typical form in the tropical
lowlands of West Colombia (from the Truando to Buenaventura),

138. PH#THORNIS SYRMATOPHORUS SYRMATOPHORUS Gould.

Phaéthornis syrmatophorus Gould, Contrib. to Ornith. 1851,
p. 139 (1851.—‘“ Interior of Quito, in Ecuador,” W. Jameson
coll.).

P. syrmatophorus Sclater & Salvin, P. Z. 8. 1879, p. 528
(Santa Elena, Medellin, Antioquia); Simon & Dalmas, Ornis,
xi. 1901, p. 217 (La Tigra, Las Cruces, Western Cordillera).

P. berlepschi K. & Cl. Hartert, Nov. Zool. i. p. 56 (1894.—Rio
Pescado, W. Ecuador).

No. 3782. @imm. La Selva, Rio Jamaraya, 4600 ft., 2.x.09.—
Wing 62; tail 75; bill 41 mm.

“Tris black, feet brown, maxilla black, mandible scarlet with
black tip.”

This bird, as well as Salmon’s skins from Medellin and Santa
Elena in the British Museum, agrees with examples from Western
Ecuador upon which Hartert’s P. berlepschi was based. I regret to
say, however, that this name is a synonym of P. s. syrmatophorus
while the eastern form has to bear the subspecific term columbianus
Boucard *, as already pointed out by Salvadori & Festa rf. Gould
established the species upon specimens obtained by Prof. Jameson.
There are three skins from this source in the Gould Collection
(now in the British Museum): one, marked “ Napo—Jameson,”

* Phaéthornis columbianus Boucard, ‘The Humming Bird, i. p. 17 (1891.—
Bogota; type in Paris Museum examined).

+ Phaéthornis syrmatophorus Salvadori & Festa, Boll. Mus. Zool. Toro, xv.
no. 868, 1900; p. 3 (Gualea, W. Ecuador). ©

1180 MR. C. E. HELLMAYR ON THE

belongs to the eastern race, P. s. columbianus, having the malar
streak and a broad stripe down the middle of the lower surface,
from the chin to the anal region, clear white, and the rump as
well as the upper tail-coverts uniform ochraceous buff; the two
others, labelled ‘“‘ Ecuador—Jameson,” agree with the so-called
“< berlepschi” from W. Kcuador, having the malar streak bright
buff, the rump feathers green with buff edges, and the whole of
the lower parts, with exception of a white stripe along the middle
of the throat, ochraceous buff. As will be easily seen from Gould’s
account, his description is a mixtum composituwm, the characters
of the lower parts (“‘ chest, abdomen, and under tail-coverts rich
buff”) being taken from the western form (berlepschi), while the
words “rump and upper tail-coverts rich buff” apply only to
the eastern colwmbianus. Since no marked type of P. syrmato-
phorus exists, M. Boucard was perfectly justified—according to
the rule of the first reviser—in restricting the name to one of
the components, and his name columbianus has to stand for the
eastern race. Thus we have :—

(a) P. s. syrmatophorus (Gould) (syn. berlepschi Hart.).
Western Heuador (Gualea, Rio Pescado, etc.) and Western
Cordillera of Colombia (La Selva, La Tigra, Las Cruces,
Medellin, Santa Elena, etc.).

(b) P. s. columbianus Boucard. Eastern Ecuador (Rio Pastaza
etc.), and Eastern Cordillera of Colombia (Bogota coll.).

139. EUTOXERES AQUILA SALVINI Gould.

[Trochilus Aquila Bourcier, P. Z. 8. 1847, p. 42 (1847—
“Jes environs de Bogota”). |

Hutoweres salvint Gould, Ann. Mag. N. H. (4) i. p. 456
(1868.—Veragua).

E. aquila (nec Bourcier) Simon & Dalmas, Ornis, xi. p. 218
(Plano de los Monos, Western Cordillera).

No. 2445. gad. Noanama, 100 ft., 5.1.08.—Wing 76; tail
54; bill 303 mm.

“Tris black, feet black, plantar surface yellow, maxilla black,
mandible yellow.”

This bird belongs to ZL. a. heterwra, as defined by Hartert &
Hartert*, having much less white in the tail than Bogota-skins
(=true LH. a. aquila). Messrs Salvadori & Festa ft cast some
doubts on the distinctness of H. baroni Hart. & Hart. t, and
take this “ species ” for the young of the ordinary heterura (sensu
Hartert). JI have neither material nor time to enter into that
question now: but whatever the specimens with an olive-grey tail
and minute, if any, white spots to the rectrices may be, there
can be no doubt that it isto this form that Gould’s term JL.
heterura has to be restricted. Although Gould, in the introductory

* Nov. Zool. i. 1894, pp, 53-54.
+ Boll. Mus. Zool. Torino, xv. no. 268, 1900, p. 2.
t Nov. Zool, i. p. 54 (1894.—Rio Pescado, Naranjal, Ecuador).

BIRDS OF WESTERN COLOMBIA, 1181

remarks to his paper, alludes to the variability of the tail-markings,
yet the formal description (J. ¢. p. 456) of H. heterura clearly
refers to the birds afterwards separated as H. baroni, cfr.
especially the words: “ tail olive-grey, 7m some instances tipped
with sullied white.”

The next available name for ZH. heterwra Hart. (nec Gould)
seems to be ZL. salvini Gould, based upon specimens from Veragua
which I cannot satisfactorily distinguish from those taken in
W. Colombia and W. Ecuador.

140. FLORISUGA MELLIVORA MELLIVORA Linn.

Trochilus mellivorus Linneus, Syst. Nat. x. p. 121 (1758—ex
Edwards: Surinam).

Florisuga mellivora Simon & Dalmas, Ornis, xi. p. 218 (Las
Cruces, West. Cordillera) ; Sclater & Salvin, P. Z. 5. 1879, p. 529
(Remedios).

Nos. 2055, 2238. Gad, gimm. Noanama: 3.ix.; Cajon,
R. Cajén : 3.x1.08.— Wing 66, 65; tail 38, 35; bill 18, 19mm.

‘Tris, feet, and bill black.”

We have also a number of skins obtained by the late J. H.
Batty on the Rio Zapota, in August and September 1898.

I’, mellivora is a wide-spread species.

14]. PoLYERATA ROSENBERGI Bouc.

Polyerata roserbergi Boucard, Genera of Humming Birds, p. 399
(end of 1895.—Rio Dagua, W. Colombia): idem, ‘ The Humming
Bird,’ v. Dec. 1895, p. 6.

Nos. 2247, 2351. ¢ Sg ad. Novita: 7, 28.xi. 08.—Wing 55,
563; tail 30, 34; bill 22, 203. mm.

No. 2551. dg ad. Juntas (Rio Tamana) : 20.11.09.— Wing 57 ;
tail 32; bill 20 mm.

No. 2570. dad. Rio Condoto: 26.111.09.—Wing 55; tail
314; bill 21 mm.

No. 2373. ¢ imm. Novita: 5.xii.08.—Wing 55; tail 32;
bill 20 mm.

No. 2255. @Q ad. Novita: 9.x1.08.— Wing 503; tail 29; bill
21 mm.

“ Tris, bill, and feet black.”

The original examples, obtained by Mr. Rosenberg in the same
district in 1894, were all more or less immature. ‘The fine series
now forwarded by Mr. Palmer enables me to state that none of
the characters which served to distinguish the Hcuadorian race
P.r. reint* holds good, and I am afraid the latter name will have
to be placed as a synonym of P. rosenbergi. Thanks to the kind-
ness of Dr. von Lorenz, of the Vienna Museum, I have been able
to examine a series of seven specimens of the so-called reinz:
266 ad,2 6 5 imm., 3 2 9, all from N.W. Ecuadort. The

* Polyerata reina Berlepsch, Orn. Monatsber. v. p. 58 (1897.—Western Ecuador).

+ They are from the following localities : Carondelet (60 ft.), Bult (60 ft.), San
Javier (60 ft.), Pambilar (60 ft.), Charco Redondo (160 ft.).

1182 MR. C, E. HELLMAYR ON THE

adult males from W. Colombia have the throat and foreneck of
exactly the some golden green hue as the Ecuadorian ones, while
the violet-blue pectoral area, in the latter, is by no means less
extended. There is no difference in the development of the
bronzy base to the outer rectrices between the two series, and the
dimensions are practically the same. The female from Novita
differs from the three “‘ reint” (ew Ecuador) by having the throat:
less spotted with shining green, though this is very insignificant.

Measurements :—
P. rosenbergi.
Wing. Tail. Bill.
mm. min. mm.
Four adult males from W. Colombia... 55-57 30-34 204-22
One immature male __,, 5 Jase DS) 32 20
One female - x as 0S 29 21

OE. ERAONOUN
Two adult males from N.W. Eeuador... 53,55 30,32 22

Two immature males __,, <s Sag 5a Oeahe Bi 21
Three adult females Be “A ... 90-52 ° 27,28 21-22

P. rosenbergi is thus found to range over the humid, tropical
coast-district of N.W. Ecuador and W. Colombia.

142. URANOMITRA FRANCLE Bourc. & Muls.

Trochilus Francie Bourcier & Mulsant, Ann. Sci. phys. et nat.,
d’ Agric. et d’Industr. Lyon, ix. p. 324 (1846.—Bogota ; descr.
¢ ad.)

Cyanomyia francie Sclater & Salvin, P. Z. 8. 1879, p. 530
(Antioquia).

Agyrtria Francie Simon & Dalmas, Ornis, xi. p. 218 (La Tigra,
Western Cordillera).

Uranomitra francie Boucard, ‘'The Humming Bird,’ vy. p. 6 (Rio
Dagua).

Nos. 2733, 2736, 3756. Gg ad. Pueblo Rico, 5200 ft., 16,
18.viii.; Siaté: 15.ix.09.—Wing 55; tail 35, 36; bill. 21-24 mm.

No. 2735. (¢)juv. Pueblo Rico: 18.viii.09.—Wing 56; tail
34 mm.

“Tris and feet black, maxilla black, mandible brown, tip black.”

Similar to Bogota-skins.

U. francie is as yet known only from the mountains of
Colombia.

143. AMAZILIA TZACATL JUCUNDA Heine.

[Zrochilus Tzacatl De la Llave, Registro Trimestre, 11. No. 5,
p. 48 (1833.—Mexico) *.]

Eranna jucunda Heine, Journ. f. Orn. xi. p. 188 (1863.— Baba-
hoyo & Esmeraldas, W. Ecuador).

* Ofr. Richmond, Auk, xvi. 1899, p. 324.

BIRDS OF WESTERN COLOMBIA. 1183

Amazilia fuscicaudata (nec Fraser) Boucard, ‘The Humming
Bird, v. p. 7 (Rio Dagua).

A. riefferi Sclater & Salvin, P. Z. S. 1879, p. 530 (Antioquia) ;
Simon & Dalmas, Ornis, xi. p. 221 (Naranjo).

Nos. 2056, 2057, 2075, 2488. ¢S gad. Noanama: 3.1x.08.;
Novita: 23.1.09; Rio Sipi: 10.ix.08— Wing 57-59; tail 35-36 ;
bill 213—22 mm.

No. 2410. @ juv. Noévita: 18.xii.08.—Wing 57; tail 34;
bill 22 mm.

“Tris and feet black, bill brown, tip black ; maxilla black in the
young bird.”

Identical in colour and dimensions with specimens from Western
Kceuador.

A. t. jucunda, which inhabits the low countries of Western
Ecuador and Western Colombia, agrees with the typical Central
American race in shape and length of the bill, but may be distin-
guished by having the upper mandible fleshy brown (instead of
blackish), and the abdomen conspicuously darker smoke-grey.

A, t. fuscicaudata, from Eastern Colombia (Bogota) and the

mountains of Merida, is much smaller, and has a considerably
shorter, stouter bill.

144, CHLOROSTILBON PUMILUS Gould.

Chlorostilbon pumilus Gould, Ann. Mag. N. H. (4) ix. p. 199
(1872.—Citado and Pallatanga, W. Ecuador); Simon & Dalmas,
Ornis, xi. p. 221 (Naranjo, La Tigra, Western Cordillera of
Colombia).

C. angustipennis (errore) Sclater & Salvin, P. Z. 8. 1879, p. 531
(Medellin).

No. 2732. oad. Pueblo Rico: 16.viii.09.—Wing and tail

in moult; bill 133 mm.

“Tris, feet, and bill black.”

This specimen is typical of C. puwmilws, which had already been
obtained in the same region by Eugéne André, as well as further
north, in Antioquia, by the late '[. K. Salmon. Topotypical birds
from Ecuador I have not seen.

(. pumilus inhabits the mountainous ranges of Ecuador and
Western Colombia.

145. THALURANIA FANNYI Del. & Boutce.

Trochilus Fannyi Delattre et Bourcier, Rev. Zool. ix. p. 310
[1846—“les bords de la rivitre de Dagua, pres Saint-Bonaven-
ture (Nouvelle-Grenade)” |.

Thalurania Fannie Simon & Dalmas, Ornis, xi. p. 221 (Buena-
ventura, El Paillon, Naranjo); Boucard, ‘The Humming Bird,’ v.
p- 6 (Rio Dagua).

No. 2814. oad. Pueblo Rico, 5200 ft., 26.x.09. Wing 55;
tail 39; bill 20 mm.

1184 MR. C. E. HELLMAYR ON THE

Nos. 2443, 3773. ¢ ¢ imm. Noanama, 100 ft., 4.1.5 Siaté,
5200 ft., 24.ix.09.—Wing 54; tail 39, 36; bill 20, 18 mm.

Nos. 2051, 2447. @ 9 ad. Noanama: 2.ix. 08, 5.1.09.—Wing
50, 51; tail 31, 322; bill 20 mm.

“ Tris and bill black; feet in males black, in females grey.”

The differences between 7’. fannyi and 7’. verticeps Gould*,
from Western Ecuador, have been well pointed out by Simon &
Dalmas. The adult male shows all the characters assigned to
Jannyi, viz , the very dark, almost black colour of the occiput and
mantle, the nearly complete, violet-blue interscapular band, the
deeply forked tail, etc. The blue margin bordering posteriorly the
glittering green crown? is not a constant feature, it being well
developed in No. 3773, scarcely indicated in No. 2443, and com-
pletely absent in No. 2814(¢ ad.). Both of the females have the
shining bluish shoulder-patch as described by Simon & Dalmas.

T. fannyi is restricted to Western Colombia, ranging from
sea-level up to about 5000 feet in the western slope of the Coast
Cordillera ¢. In Western Ecuador it is replaced by 7’. verticeps
Gould, which may be only subspecifically separable.

146. CHALYBURA UROCHRYSA Gould.
Hypuroptila urochrysa Gould, P. Z. 8. 1861, p. 198 (1861.—

«« Panama ”’).

Chalybura buffoni (errore) Sclater & Salvin, P. Z. 8. 1879,
p. 529 (Remedios, Santa Elena).

H. urochrysea Salvin, Cat. B. Brit. Mus. xvi. p. 89 (Medellin,
Remedios).

Chalybura urochrysea Hartert, Nov. Zool. v. 1898, p. 494
(Cachabi, N.W. Ecuador).

Nos. 2083, 2329, 2581. ¢ gd ad. Sipi: 12.ix.08; Ndévita:
23.x1.08 ; Condoto: 1.iv.09.—Wing 67-704; tail 42-46; bill
233-24 mm.

No. 2218. dimm. Rio Cajon: 28.x.08.— Wing 684 ; tail 42 ;
bill 24 mm.

Noss eZ 091 20905) 2b 22.2). OVO wads Ye SipitlSre 2 pipe.
6.x.08; Rio Cajon : 29.x.08.— Wing 62-66 ; tail 38-42; bill 214-
24 mm.

Tris black, feet pink, maxilla black, mandible pink (@ ), light
brown (¢).”

This rare species is easily recognizable by its bronze-green tail,
combined with the pale mandible. The females have the six
outer tail-feathers distinctly tipped with greyish white, and
the under tail-coverts pale mouse-grey (brownish grey). The
specimens obtained by Salmon in Antioquia I have examined

* In Jardine’s Contrib. to Ornith. 1851, pt. ii. p. 79, pl. xxi. (Apr. 1851.—Quito,
Ecuador),

+ Cfr. Hartert, Tierreich, livr. 9, 1900, p. 85.

i The reported occurrence at Popayan and Pasto (!) does not rest upon reliable
authority.

BIRDS OF WESTERN COLOMBIA. 1185

in the British Museum, and found them to belong to the present
species. Strangely enough, they had been determined by Salvin as
C. buffoni.

C. urochrysa is, according to our actual knowledge, confined to
the dense forests along the Pacific coast of N.W. Ecuador and
W. Colombia *. It is particularly common in the lowlands,
though Salmon also procured examples near Remedios, at 2300
feet elevation.

At higher altitudes, however, in the Western Cordillera another
species of Chalybura occurs, specimens of which were obtained
at Rio Dagua t by Rosenberg, and at Naranjét by E. André.
It agrees with C. buffoni from Bogota, in the coloration of the
tail (rectrices bluish black, more or less edged with bronze green)
and lower parts, but has the lower mandible mostly pale
brownish, like C’. wrochrysa. With only a single example before
me I refrain, however, from naming this apparently undescribed
form.

147. BoISSONNEAUA FLAVESCENS FLAVESCENS Lodd.

Trochilus flavescens Loddiges, P. Z.8. 1832, p. 7 (1832.—
Popayan, Colombia).

Panoplites flavescens Sclater & Salvin, P. Z. S. 1879, p. 529
(Medellin).

No. 2793. gad. Tatamd Mountain, 8000 ft., 12.x.08.—Wing 80;
tail 52; bill 174 mm.

“ Tris, feet, and bill black.”

This bird fully agrees with Bogota skins except that the green
tip to the inner web of the outermost rectrix is a trifle longer.
Keuadorian specimens have much broader green tips to the lateral
rectrices, and the buff portions of the tail as well as the under
tail-coverts are deeper in tint. They have been separated by Mr.
Oberholser as L. f. tinochlora §.

148. HRIocNEMIS AURELIZ Bourc. & Muls.

Trochilus Aurelie Bourcier & Mulsant, Ann. Sci. phys. et
nat., d’Agric. et d’Industr. Lyon, ix. p. 315 (1846.—Bogoti).

Eriocnemis aurelie Sclater & Salvin, P. Z. S. 1879, p. 530
(Santa Elena, Medellin).

No. 2737. 3 juv. Pueblo Rico, 5200 ft.: 6.ix.08.—Wing 63 ;
tail 40; bill 19 mm.
“Tris black, feet grey, bill black.”

Notwithstanding its rather long bill, this specimen appears

* The original locality “ Panama” is open to doubt. Warscewicz was very
careless about labelling, and, moreover, not one of the many later travellers met
with the species on the Isthmus.

+ Hypuroptila buffoni Boucard, ‘The Humming Bird,’ v. 1895, p. 8 (specimen
examined by me).

t Chalybura buffoni Simon & Dalmas, Ornis, xi. p. 222 (Naranjé, 1900 ft.).

§ Proc. U.S. Mus. xxiv. p. 329 (1902.—West side of Corazdn, Ecuador).

1186 MR. C. E. HELLMAYR ON ‘THE

to belong to the present species, for in coloration it closely
resembles several Bogota skins in corresponding stage.

E. aurelic is well-known as an inhabitant of the mountains of
Colombia.

149. UrostrictE BENJAMINI Boure.

Trochilus Benjamini Bourcier, Compt. Rend. Ac. Sci. Paris,
xxxi. no. 6, Febr. p. 187 (1851—“les régions chauds des
environs de Gualea,” W. Ecuador).

No. —(¢) ad. la Selva, Rio Jamaraya, 4600 ft., Oct. 09.—
Wing 52; tail 35; bill 23 mm.

Compared with specimens from Western Ecuador this bird has
a decidedly longer bill and more white along the middle of the
belly. From a single skin it is difficult to say whether these
divergencies are of any importance.

Like ordinary benjamini the specimen from La Selva has a
large lilac prepectoral spot.

So far as I am aware this is the first Colombian record of
U. benjamini, although it is fairly common in Ecuador on both
sides of the Andes.

150. HELIANGELUS ExoRTIS Fras.

Trochilus exortis Fraser, P. Z. S. 1840, p. 14 (1840.—Guaduas,
Colombia).

Heliotrypha parzudakii Sclater & Salvin, P. Z. S. 1879, p. 529
(Santa Elena). aed

No. 2792. g ad. Tatama Mountain, 8000 ft., 12.x.09.—
Wing 69; tail 49; bill 17 mm.
“ Tris, feet, and bill black.”

A common bird in the mountains of Colombia and northern
Ecuador. Palmer’s specimen agrees well with Bogoti skins.

151. Heuiorurix BArroti Boure. & Muls.

Trochilus Barroti Bourcier et Mulsant, Ann. Sci. phys. et nat.,
d’Agric, etc. Lyon, vi. p. 48 (1843.—* Carthagéne,” Colombia).

Heliothri« barroti Sclater & Salvin, P. Z. 8. 1879, p. 529
(Remedios).

No. 2198. g ad. Noanama; 17.x.08.—Wing 66; tail 42;
bill 16 mm. —

No, 2329. g¢ juv. Névita; 23.xi.08—Wing 65; tail 45 <
bill 154 mm.

“Tris, feet, and bill black.”

Not apprecially different from Costa Rican skins. Z. barroti,

which is very likely only a geographical race of H. auritus,
ranges southwards as far as Western Ecuador,

BIRDS OF WESTERN COLOMBIA. 1187

152. CYANOLESBIA KINGII, subsp.

[Ornismya king Lesson, Hist. Nat. Trochil. p. 107, pl. 38
(1832. —“* Jamaique,” errore — description and plate evidently
apply to the common blue-tailed Bogota form, with a large violet-
blue gular patch)].

No. 3784. gad. Tatama Mountain, 4600 ft. 7.x.09.— Wing 74;
tail (slightly moulting) 93; bill 14 mm.

“Tris, feet, and bill black.”

This bird cannot be identified with any of the known forms.
It is somewhat intermediate between OC. k. kingii from Bogota,
and C. k. cwlestis * from Ecuador, though apparently different
from either. With the latter, it shares the long, robust beak, the
broad, rufous-buff edges to the under tail-coverts, and the bronzy-
green general tone of the lower parts; but the rufescent margins
to the feathers of the breast and abdomen are completely absent,
and the colour of the tail is more like Bogoté specimens. It
should be mentioned, however, that the under surface of the
rectrices is much duller, more blackish than in any other example
examined by me. More material will probably show the West
Colombian birds to constitute a distinct race. Curiously enough,
both Hartert 7, and Simon & Dalmas{ record the green-tailed
C. k.emme Berl., without gular spot, from the Western Cordillera,
while the Tatama bird has the throat extensively violet-blue.

153. CHLORONERPES RUBIGINOSUS GULARIS Harg.

[Picus rubiginosus Swainson, Zool. Ilustr. i. pl. xiv. (1820-1.—
“Spanish Main,” 7. e. Venezuela, environs of Cumans) |].

Chloronerpes gularis Hargitt, Ibis, (6) i. p. 230 (1889.—Santa
EKlena, Antioquia).

C. rubiginosus (errore) Sclater & Salvin, P. Z. S. 1879, p. 533
(Retiro, Concordia, Santa Elena).

_ No. 2798. g ad. Loma Hermosa, Rio Jamaraya, 4150 ft.,

18.x.09.— Wing 122; tail 84; bili 263 mm.

“ Tris dark brown, feet plumbeous, bill black.”

This bird agrees perfectly with Hargitt’s description. It differs
from C. 7. rubiginosus, from N.E. Venezuela (Cumana) and
Trinidad, in having the rump and tail-coverts pale yellow in
decided contrast to the deep tawny olive of the upper back, and
the under parts more richly yellow with the blackish bars
conspicuously narrower and confined to the foreneck and breast:
The throat is much less variegated with whitish, and the feathers
of the pileum, from the forehead to the nape, are tipped with
crimson.

C. r. gularis is limited to Western Colombia, inhabiting both
the Coast and the Central Cordillera.

* Cynanthus coelestis Gould, Introd. Humming-Birds p, 102 (1861.—Ecuador).
+ Tierreich, Lief. 9, 1900, p. 176.
{ Ornis, xi. 1901, p. 223 (Las Cruces, W. Cordillera),

1188 MR. C. E. HELLMAYR ON THE

154. MELANERPES PUCHERANI PUCHERANI Malh.

Zebrapicus pucherani Malherbe, Rev. Mag. Zool. (2) i. p. 542
(1849.—“ Tabago,” errore.*).

Nos. 2258, 2279. ¢g dad. Névita: 10, 13.xi.08.— Wing 115,
113; tail 67, 65; bill 26, 25 mm.

Nos. 1970, 2259. 9 2 Guineo: 5.vui.; Névita: 10.xi.08.—
Wing 108; tail 60, 62; bill 21 mm.

‘“‘ Tris brown, feet grey or greyish green, bill black.”

The series agrees with specimens from Western Ecuador.

M. p. pucherani ranges from Nicaragua southwards through
Western Colombia to Guayaquil, S.W. Ecuador. Birds from
Central America are perhaps slightly different, but I have not
sufficient material to decide the question. The form occurring
from Honduras north to 8. Mexico has lately been separated as
M. p. perileucus Todd 7.

155. VENILIORNIS KIRKII CECILIIT Malh.

[Picus (Chloropicus) Kirkii Malherbe, Rev. Zool. viii. p. 400
(1845.—Tobago)].

Mesopicos Cecilii Malherbe, Rev. Mag. Zool. (2) i. p. 538
(1849.—Colombia).

Chloronerpes cecilie Sclater & Salvin, P. Z. 8. 1879, p. 533
(Antioquia, Remedios, Neche).

No. 1986.( 3.) ad. Guineo, Rio Calima : 10.viii.08.— Wing 84 ;
tail 55; bill 201 mm.

ub Th brown, feet pinkish grey, maxilla dark grey, mandible
paler.”

This bird, in abraded, bleached summer plumage, is not
appreciably different from Bogotaskins. Specimens from Western
Ecuador also agree well with the latter.

V. k. cecilii is peculiar to Colombia (Coast, Central and Kastern
Cordillera £) and Western Ecuador. In Panama it is replaced
by the doubtfully separable V. k. darienensis Ridgw. §

156. CrLeus LoricAtus Reichb.

Meiglyptes loricatus Reichenbach, Scansorie, p. 405, pl. delxxxi.
figs. 4495-96 (1854.—“ Peru”; descr. 9 ).

Celeus loricatus Sclater & Salvin, P. Z. S. 1879, p. 533
(Remedios & Neche).

* The species does not occur on “ Tabago”’ (sc. Tobago), the original locality being
no doubt erroneous. Later, in the Monogr. des Picid. ii. p. 227, Malherbesays that
the specimens he saw in the British Museum came from “ Tabago ” while those
existing in his own collection were from “ Nouvelle-Grenade.” We may therefore
accept Colombia as the type locality since the birds from this country agree very
well with Malherbe’s description and figure (pl. ciii.).

+ Proc. Biol. Soc. Wash. xxiii. p. 154 (1910.—Manatee, British Honduras).

~ Count Berlepsch (J. f. Orn. 1884, p. 314: Chloronerpes cecilie) notices that
examples from Bucaramanga are considerably larger than those from Bogota.

§ Proc. Biol. Soc. Wash. xxiv. p. 33 (1911.—E] Réal, Darien).

BIRDS OF WESLERN COLOMBIA. 1189

C’. mentalis Cassin, Proc. Acad. N. Sci. Philad. 1860 p. 137
(1860.—Turbo & R. Atrato; deser. 3).

No. 2254. (¢) ad. Noévita: 9.x1.08.—Wing 117; tail 70;
bill 21 mm.

‘Tris crimson, feet grey, maxilla black, mandible yellow.”

This specimen, a fully adult male with cheeks, malar region
and upper throat crimson, corresponds exactly with Cassin’s
description of C. mentalis. ‘The types of the latter were obtained
in the same general region, but farther to the north, on the Rio
Atrato.

It appears, however, that J/. loricatus was based upon a female
of the same species,

C. loricatus ranges from Costa Rica to Western Ecuador.
I have not seen specimens from Costa Rica or Panama. If
separable, they will have to bear the name squamatus Lawr. *

157. CEOPHLG@US LINEATUS LINEATUS Linn.

Picus lineatus Linneus, Syst. Nat. 12, 1. p. 174 (1766—ex
Brisson : Cayana).

Dryocopus lineatus Sclater & Salvin, P. Z. 8. 1879, p. 532
(Santa Elena).

No. 2014. 9 ad. Noanama: 25.viu1.08.— Wing 180; tail‘115;
bill 36 mm.

“‘ Tris white, feet blue-grey, bill black.”

[158. PicUMNUS OLIVACEUS GRANADENSIS Lafr.

[Picumnus olivaceus Lafresnaye, Rey. Zool. vill. p. 7 (1845—
“ad. Bogotam, in Colombia ”). |

PR: granadensis Lafresnay Cli nice x. ype 1S. iC S4i ad:
Caly, in Nova Granada ”—coll. Delattre: @ ad., type in Mus.
Acad. Philad., cfr. Stone, Proc. Acad. N. Sci. Philad. li. 1899,
p- 52) ; Hargitt, Cat. B. Brit. Mus. xviii. p. 549 (part.: fA g.
¢ 2 Medellin, Antioquia, examined by me).

P. olivaceus (nec Lafr.) Sclater & Salvin, P. Z. 8. 1879, p. 532
(Medellin, Colombia).

P. canus Bangs, Proc. Biol. Soc. Wash. xxi. p. 72 (1910.—
Naranjito, R. Dagua; one @ ad.).

No. 387. g ad. Primavera, W. Colombia, 5200 ft. Raap coll.—
Wing 60; tail 30; bill 133 mm.

No, 246. ¢ ad. San Isidro, 2700 ft. Raap coll.—Wing 59;
tail 29% bill 12) mm.

Nos. 257, 271.
Raap coll. Wing 58, 59; tail 29, 30; bill 12 mm.

One 2 ad. San Antonio, Western Cordillera, 5400 ft. 13.vi.09.
Fassl coll.— Wing 57; tail 30; bill 12 mm.

Two 6 g ad. Rio Dagua: 12, 16.vi.95. Rosenberg coll.—
Wing 58; tail 292, 30; bill 12-13 mm.

2700 ft.

* Celeus squamatus Lawrence, Ibis, v. p. 184 (1863.—Panama Railroad).
Proc. Zoou. Soc.— 1911, No. LX XX. 80

1190 MR. C. E. HELLMAYR ON THE

This series, which belongs to the Tring Museum, enables us
at last to make out the status of P. granadensis, originally based
upon an adult female from the same district (Cali). I am sorry
to say that Mr. Bangs under P. canus simply redescribed the
true granadensis. This is quite evident on comparing the two
descriptions, which agree almost word for word. In a note
appended to the diagnosis of P. o. harterti *, I have already
alluded to the distinguishing characters of P. 0. granadensis, but
a few additional remarks may not be out of place here. The
geographical variation of this group was quite unsatisfactorily
understood until Mr. Hartert ~ pointed out the differences
existing between the three races then known to him. MHartert
was, however, not acquainted with granadensis Lafr., for the
birds from Western Ecuador to which he applied that name proved
to be distinct and were subsequently separated by me as P. o.
hartertt.

According to our present knowledge, the following subspecies
of P. olivaceus ave easily recognizable :—

(a) P. otrvacnus oxivAceus Lafr.

Type from Bogota.

Adult. Back warm greenish brown or olivaceus brown; chest
strongly tinged with deep olivaceous; breast and abdomen
yellowish with well-defined, though narrow, dusky stripes,
particularly on the lower breast and flanks; quills edged with
olivaceous. Adult males with tips to feathers of forehead
“flame scarlet” (vather duller than pl. vu. fig. 14 of Ridgway’s
Nomenclature).

Hab. Appavently the Eastern Cordillera of Colombia. All the
specimens I have seen, eleven at Tring, and nine in the Munich
Museum, are of the well-known Bogota form.

(6) P. oLIvaceus FLAVorINcTUS Ridgw.t

Type from Pozo Azul, Western Costa Rica.

Adwt. Back not unlike P. 0. olivaceus, but mostly somewhat
darker ; edges of remiges brighter and deeper olivaceous yellow ;
chest decidedly darker, more brownish, less greenish; dusky
stripes of breast and abdomen wider and less sharply defined.
Adult males with tips to feathers of forehead dull “cadmium
orange” (Ridgw. Nomencel. pl. vi. fig. 2.)

iah South-western Costa Rica (south of Pozo Azul de Pirris)
and Chiriqui §.

* Bull. B. O. C. xxiii. March 1909, p. 67.

+ Novit. Zool. ix. 1902, pp. 606-7.

t P. flavotinctus Ridgway, Proc. U.S. Mus. xi. 1888, p. 543 (1889.—Pozo Azul,
vw pe Rica).

§ An additional. subspecies, P. clivaceus panamensis Ridgw. (Proc. Biol. Soc.

W Hae xxiv. 1911, p. 44: Lion Hill Station, Panama) inhabits Panama (Lion Hill,
Tocoumé). There is a single adult male trem the latter locality, obtained
by E. André, in the Tring Museum. This form is nearest to P. o. flavotinctus, but

has the chest buffy yellow (instead of brownish) and the back much paler, light
earthy-brown.

BIRDS OF WESTERN COLOMBIA. 1191

Examined: Two males and two females, Costa Rica (Tring
Museum), four males and one female, Chiriqui (Tring and
Munich Museums).

N.B.—An adult male in bad condition, said to be from San
Esteban, Venezuela, in the British Museum differs merely in
having the breast more strongly tinged with brown. The locality
is most probably erroneous.

(c) P. onivaceus HARTERTI Hellm.

P. o. harterti Hellmayr, Bull. B.O.C. xxiii, p. 67 (March
1909.—Paramba, N.W. Ecuador).

P. granadensis (nec Lafr.) Sclater, P. ZS. 1860, p. 95
(Nanegal) ; idem, l.c. p. 287 (Babahoyo); Berlepsch & Taczan-
owski, 1. c. 1883, p. 570 (Chimbo, Yaguachi); iidem, |. c. 1885,
p. 106 (Yaguachi) ; Hartert, Nov. Zool. v. 1898, p. 497 (Chimbo) ;
Salvadori & Festa, Boll. Mus. Zool. Torino, xv. no. 368, 1900,
p. 16 (Vinces) ; Goodfellow, Ibis, 1902, p. 209 (Santo Domingo) ;
Hargitt, Cat. B. Brit. Mus. xviii. p. 549 (part.: 7, k, Babahoyo ;
l, Monj1).

P. olivaceus granadensis Hartert, Nov. Zool. ix. 1902, p. 606
(Chimbo, Paramba, 8. Domingo).

Type from Paramba, N.W. Kcuador, 3500 ft. alt.

Adult. Much like P. o. flavotincéus, but chest olivaceous as in
P. o. olivaceus, and adult males with tips of feathers on forehead
much paler, cadmium yellow (Ridgw, pl. vi. fig. 6).

Hab. Western Ecuador, from sea-level up to 3500 feet.

Exanuned: Kleven males and four females in British, Tring,
and Munich Museums.

(d) P. oLIVACEUS GRANADENSIS Lafr.
Synonymy, see above.

Type from Cali, W. Colembia.

Adult. Differs at once from the three preceding races in having
the back light olive-grey or pale smoky-grey (without greenish
or brownish suffusion) and the under parts creamy-white, nearly
uniform, with but a few obsolete dusky streaks on the flanks, and
the foreneck very slightly tinged with dull greyish. Edges of the
quills creamy white, instead of olivaceous or olive-yellow. Tips
of feathers on forehead in adult males even paler than in
P. 0. harterti, “ chrome-yellow ” (Ridgw. pl. vi. fig. 8.

Hab. Western Cordillera of Colombia: Cali (Delatire), Rio
Dagua (Rosenberg), Naranjito (Palmer); Primavera, San Isidro,
Media Luna (aap); San Antonio (fassl); Medellin, Antioquia
(Salmon).

Obs. In addition to the specimens enumerated above I
examined a couple from Medellin (Salmon) in the British Museum
and found them toagree. One of the Dagua examples (Rosenberg)
is rather more brownish grey above than the others, thereby
forming the passage to P. 0. harterti.

80*

1192 MR. C. E. HELLMAYR O THE

(e) P. oLtvAcEUS DIMotUS Bangs.

Picumnus dimotus Bangs, Bull. Mus. Harvard Coll. xxxix.
p. 146 (1903.—Ceiba, Honduras).

P. olivaceus (nec Lafr.) Hargitt, Cat. B. Brit. Mus. xviii. p. 548
(part.: A, 7, Julian, S. Pedro: Honduras).

Adult. Much like P. o. olivaceus, and agreeing in the adult males
in having the tips of the feathers on the forehead scarlet, but with
back and chest much more greenish, and dusky streaks on belly
less distinct.

Hab. Honduras: Ceiba, Julian, San Pedro; Nicaragua.

Kxamined : Two adult males from Honduras (Whitely) in the
British Museum. |

159. CERYLE TORQUATA TORQUATA Linn.

Alcedo torquata Linneeus, Syst. Nat. 12, i. p. 180 (1766—ex
Brisson: Mexico & Martinique).

Ceryle torquata Cassin, Proc. Acad. N. Sci. Philad. 1860, p. 133
(R. Atrato & Truando); Sclater & Salvin, P.Z.S. 1879, p. 534
(Neche). !

No. 3759. Q ad. Siaté, R. Siaté, 5200 ft., 17.1x.09.—Wing
195; tail 122; bill 65 mm.

“Tris brown, feet greyish green, bill black, base yellow.”

160. CERYLE AMAZONA Lath.

Alcedo amazona Latham, Ind. Ornith. i. p. 257 (1790.—
“ Cayana ”).

Ceryle amazona Cassin, Proc. Acad. N. Sci. Philad. 1860, p. 133
(R. Nercua); Sclater & Salvin, P. Z.8. 1879, p. 534 (Neche).

No. 2397. gad. Ndéovita: 14.x11.08.—Wing 131; tail 85;
bill 66 mm.

‘“‘ Tris dark brown, feet and bill black.”

161. CeryLe inpA Linn.

Alcedo inda Linneus, Syst. Nat. 12, 1 p. 179 (1766—
ex Edwards: ‘“ India occid.”, errore; we substitute Surinam
as type locality).

Ceryle inda Cassin, |. ce. p. 133 (Turbo).

No. 2723. Q ad. ‘Tadd, R. San Juan: 28.11.09.—Wing 97;
tail 62; bill 43 mm.

“Tris dark brown, feet black, maxilla black mandible dark
brown.”

162. CERYLE AMERICANA AMERICANA Gm. (?)

Alcedo americana Gmelin, Syst. Nat. 1, 1. p. 451 (1788—
ex Daubenton, Pl. Knl. 591. figs. 1, 2: Cayenne).

Ceryle cabanisi (nec Tschudi) Sclater & Salvin, P. Z.S. 1879,
p. 534 (Retiro, Concordia, Medellin).

Nos. 1990, 2077, 2209. g g ad. Mouth of Calima: 13.viii. ;
Sipi: 1l.ix.08; Noanama: 21.x.08.—Wing 75-77; tail 54-55 ;
bill 36-39 mim.

BIRDS OF WESTERN COLOMBIA. 1193

“Tris dark brown, feet and bill black.”

These three as well as another adult male from the Cauca
River (Batty coll.) in the Munich Museum differ from a very
large series of skins obtained in Cayenne, Venezuela (Caura),
Brazil, etc., in having the under tail- coverts whoily unspotted
white or very faintly spotted with bronze-green. It should be
mentioned, however, that this character is likewise found in two
specimens from the island of Toba go.

In the absence of typical examples from Western Peru, 1
cannot say whether the birds from Western Colombia should not
be referred to C. a. cabanisii Tsch.* I notice, however, that
the white bars on the quills are by no means broader than in
typical C. a. americana, which is said to be the case in the
Peruvian form.

163. TROGON MASSENA Gould.

Trogon massena Gould, Monogr. Trogon. Ist ed. pl. 16 (1838.—
Guatemala); Cassin, Proc. Acad. N. Sci. Philad. 1860, p. 135
(Truando ; delta of the Atrato).

No. 2008. g ad. Noanama, 100 ft., 26.viii.08.—Wing 170;
tail 168; bill 24 mm.

“Tris dark brown, feet brownish yellow, bill orange-yellow.”

Compared with others from Central America, this bird is
smaller, and has the middle pair of rectrices washed with dull
bluish instead of clear bronze-green. With a single specimen at
hand it is, of course, impossible to judge the value of these
variations. In other respects the male from Noanama is a
typical massena, there being not the least trace of a white
breast-band, ete.

Noanama is the most southerly locality yet known for 7’.
massena. Curiously enough, the late T. K. Salmon obtained
the nearly allied 7. melanurus macrowrus Gould in the State
of Antioquia T, provided that his specimens have been correctly
determined.

164. UrosPaArHA MARTI SEMIRUFA Ncl.

[Prionites martw Spix, Av. Bras. i. p. 64, pl. Ix. (1824—“ in
sylvis Parae”).|

Momotus semirufa Sclater, Rev. Mag. Zool. (2) v. p. 489
(1853—part.: Santa Marta, New Grenada {).

Momotus Martii (nec Spix) Cassin, Proc. Acad. N. Sci. Philad.
1860, p. 1386 (River Nercua).

Urospatha mariwu Sclater & Salvin, P. ZS. 1879, p. 534
(Remedios, Neche).

* Alcedo Cabanisii Tschudi, Fauna Peru., Aves, p. 253 (1844-45.—“ environs of
Lima’”’).

+ T. macrurus Sclater & Salvin, P. ZS. 1879, p. 535 (Remedios, Neche).

{ Although one of the localities, Rio Javarri (W. Brazil), refers to typical U. m.
inane, Sclater’s description Is evidently taken from the Santa Marta specimen.
Cfr. the words: “ cauda spatulata.”

1194 MR. C. E. HELLMAYR ON THE

Nos. 2108, 2419. ¢ gad. Sipi: 23.ix.; Névita: 20.xii.08.—
Wing 147, 150; tail 250, 255; bill45, 47 mm.

No. 2013. 9 ad. Noanama: 25.viii.08.—Wing 148; tail 250;
bill 46 mm.

“Tris dark brown, feet and bill black.”

All three specimens have the median rectrices spatulated and
decidedly bluish. Cfr. also my remarks in Abhandl. Bayer.
Akad. Wiss., II. Cl. vol. xxii. 3, 1906, p. 611, and in Nov. Zool.
xiv. 1907, p. 403.

U. marti semirufa replaces the typical race in Western
Keuador and Colombia, ranging northwards to Costa Rica and
Nicaragua.

165. Momotus 2QUATORIALIS ZQUATORIALIS Gould.

Momotus equatorialis Gould, P. Z. 8. 1857, p. 223 (Jan. 1858.—
‘“‘ Archidona, near the Equatorial line, on a branch of the Rio
Napo”); Sclater & Salvin, P.Z.S. 1879, p. 534 (Envigado,
Retiro, Concordia, Frontino).

No. 2820. g ad. Pueblo Rico, 5200 ft., 29.x.09.—Wing 154;
tail 315; bill 422 mm.

‘* Tris vermilion, feet dark grey, bill black.”

We have also a fine pair procured by the late J. H. Batty
at Rio Lima, 4500 feet, Western Cordillera, in August 1898.
The Colombian skins agree exactly with others from Ambato,
HE. Ecuador, which can be regarded as topotypical.

AM. equatorialis is an inhabitant of the highlands of Ecuador
and Western Colombia. It may prove to be merely the southern
form of the Central American M, lessoni Less. In Peru it is

replaced by a nearly allied race, J/. wquatorialis chlorolemus
Berl. & Stolzm.*.

166. GALBULA MELANOGENTA Scl.

Galbula melanogenia Sclater, Contrib. to Ornith. for 1852, p. 61
(1852—loe. ign.).

G. ruficouda (nee Cuvier) Cassin, Proc. Acad. N. Sci. Philad.
1860, p. 134 (River Nercua).

Nos. 2219, 2513. gg ad, Rio Cajén, 125 ft., 29.x.08 ;
El Tigre, 320 ft., 5.11.09.— Wing 79, 82; tail 93, 94; bill 47 mm.

Nos. 2220, 2567. 9 9 ad. Rio Cajén: 29.x.08; Juntas,
400 ft., 11.1.09.-Wing 79, 81; tail 93; bill 44 mm.

“Tris dark brown, feet yellowish green, bill black.”

The specimens are identical with others from Chiriqui and
Western Ecuador.

167. Bucco prcrorauis Gray.

Bucco pectoralis Gray, Genera Birds, i, pl. xxvi. (Dec. 1846—
no locality given); Wyatt, Ibis, 1871, p. 374 (Magdalena Valley,
between Naranjo and the river); Sclater & Salvin, P. Z.S. 1879,

* P.Z.S. 1902, ii. p. 35 (1902.—Ocobamba near Cuzco, S.E. Peru).

BIRDS OF WESTERN COLOMBIA. 1195

p- 536 (Neche); Berlepsch, Journ. f. Ornith. 1884, p. 315
(Bucaramanga).

Nos. 2464, 2626. ¢ ad, d imm. Noévita: 4.xi1.08; Tado:
29.iv.09.—Wing 100; tail 83; bill 323, 32 mm.

Nos. 2318, 2359, 2464. 9 9 ad., 1 Q imm. Novita: 20.x1.,
1.xii.08; Noanama: 12.1.09.—Wing 95-100; tail 80-84; bill
30-33 mm.

“Tris dark red, feet grey, bill black.”

Adults have the edges of the scapulars, wing-coverts, rump, etc.,
pure white, while they are more or less tinged with buff in
immature birds; the latter also show a slight buffish wash on the
lower flanks and under tail-coverts.

Two skins from Chiriqui have the black portions of the
plumage faintly glossed with oil-green, while our Chocé series,
as also a couple from N.W. Hcuador, show a strong metallic blue

loss.
B. pectoralis ranges from Western Panama (Chiriqui) south-
wards to N.W. Ecuador (province Hsmeraldas *). It is exclusively
found in the hot, dense forests of the lowlands.

168. Bucco TEcTUS SUBTECTUS Scl.

[Bueco tectus Boddaert, Tabl. Pl. Enl. p. 43 (1783—ex
Daubenton, Pl. Enl. 688. fig. 2: Cayenne). |

Bucco subtectus Sclater, P.Z.S8. 1860, p. 296 (1860.—Esme-
raldas, N.W. Ecuador); Sclater & Salvin, P.Z.S. 1879, p. 536
(Neche).

Nos. 2634, 2703. 9 9 ad. Tadoé (230 ft.): 3.v., 12.vi.08.—
Wing 69; tail 56; bill 21, 22 mm.

‘“‘ Tris red, feet and bill black.”

This well-characterized race differs from B. ¢. tectus, of Cayenne,
Para, and Hastern Venezuela (Caura), in several important points :
The upper parts are of a much deeper, more glossy black; the
white spots on the head are confined to the forehead and
anterior crown; the white cross-bar on the inner web of the
submedian pair of rectrices is either wholly absent or but faintly
indicated; the black crescent of the foreneck is much narrower ;
the bill more slender.

Birds from Panama have even less white spotting about the
forehead than those from more southern localities, but otherwise
they do not differ.

B. t. subtectus has exactly the same range as the preceding
species, occurring from Western Panama (Chiriqui) south to
N.W. Hcuador (province Esmeraldas).

169. Bucco NoANAM# Hellm.

Bucco noaname Hellmayr, Bull. B.O.C. xxv. p. 21 (1909.—
Noanama, W. Colombia).
* There are two specimens in the Tring Museum: gad. from Carondelet (60 ft.),

Oct. 16, 1900; 9 ad. Bultin (160 ft.), Dec. 23, 1900, both secured by Mr. G.
Flemming, one of Mr, Rosenberg’s correspondents.

1196 MR. C. E. HELLMAYR ON THE

No. 2033. g ad. Noanama (100 ft.), 28.vili.08. Type of
species.— Wing 81; tail 66; bill 28 mm.

Nos, 2708, 2711. § Q ad. Tado (230 ft.), 8, 16.vi.09.— Wing
81, 80; tail 66; bill 28, 27 mm.

“Tris red or brown, feet grey, bill black.”

Adult (sexes alike). Upper part of the head and nape dark
ashy; back and upper wing-coverts dark sepia-brown, each
feather with a distinct, pale rufescent apical margin ; upper tail-
coverts with rather brighter rufescent cross-bands and edges ;
remiges and rectrices dusky, narrowly fringed with rufescent-buff
along the outer web. Narrow frontal edge and broad super-
ciliaries, reaching as far as the posterior angle of the eye, soiled
white; Jores ashy black; cheeks, ear-coverts, and sides of the
neck dark ashy, some of the feathers edged with pale greyish.
Malar region, chin, aud throat white; chest dull sooty black,
most of the feathers showing, on their concealed basal portion,
a white mesial spot or streak, only to be seen when the feathers
are raised; rest of the belly white, washed with ochreous-buff
and marked with coarse blackish spots or transverse bands; under
tail-coverts uniform buff. Axillaries and under wing-coverts deep
buff, mixed with blackish ; inner web of remiges broadly edged
with buff on its basal half.

This new Puff-bird is not very nearly related to any other
member of the genus. In proportions and style of coloration
(such as the rufescent cross-bands on the upper parts, buff
axillaries, under wing-coverts and quill-lining, ete.) it is not un-
like B. macrodactylus Spix from Amazonia; but the ashy cap,
the much broader blackish breast-band, the coarse spotting of the
belly, ete., as well.as the much larger size, serve to distinguish it
at a glance.

The three specimens sent by Mr. Palmer are perfectly alike,
the female being but very slightly smaller.

B. nodname is, as yet, known only from the valley of the San
Juan River in Western Colombia.

170. MALACOPTILA PANAMENSIS POLIOPIS Scl.

[Malacopiila Panamensis Lafresnaye, Rev. Zool. x. p. 79
(1847.— Panama). |

Malacoptila poliopis Sclater, P. Z. 8. 1862, p. 86, pl. viii.
(1862.—Esmeraldas, N.W. Ecuador, descr. 9 ).

M. panamensis (nec Lafr.) Cassin, Proc. Acad. N. Sci. Philad.
1860, p. 134 (Rio Truando).

Nos. 2085, 2437, 2520. g¢ g ad. Sipi (150 ft.), 12.ix.; Névita
(150 ft.), 26.x11.08 ; El Tigre (320 ft.), 9.11.09.— Wing’ 88-90 ;
tail 75-78; bill 29 mm.

Nos. 2084, 2593. 9 @ ad. Sipi (150 ft.), 12.ix.08 ; Condoto
(150 ft.), 12.iv.09.— Wing 86-88 ; tail 75, 79; bill 29 mm.

“Iris red, feet grey, maxilla black, mandible yellowish grey.”

The series agrees perfectly with topotypical examples from the

BIRDS OF WESTERN COLGMBIA. 1197

province of Esmeraldas. Jf. p. poliopis appears to be a well-
marked race, differing from J. p. panamensis, of Panama,
Chiriqui, and 8.W. Costa Rica, in its darker coloration. The
males are much darker rufous above and have the throat
and foreneck of a deeper cinnamon-rufous shade, whilst the
females may also be distinguished by their fuscous (less brownish)
back, more blackish cheeks, darker ochraceous foreneck, etc.

M. p. poliopis is apparently confined to the lowlands and foot-
hills of the Pacific coast district of Colombia (from the Truando
southwards) and W. Ecuador (south to Guayaquil).

171. MALAcoprinaA MysTACGALIS Lafr.

Monasa mystacalis Lafresnaye, Rev. Mag. Zool. (2) 11. p. 215
(1850.—Colom bia).

Malacoptila panamensis (errore) Sclater & Salvin, P. Z.5.
1879, p. 5386 (Remedios).

M. mystacalis Berlepsch, Journ. f. Ornith. 1884, p. 315
(Bucaramanga ; crit.).

Nos. 2803, 3764. g 2 ad. Loma Hermosa (4150 ft.), 22.x. ;
Siat6, near Pueblo Rico (5200 ft.), 21.1x.09.— Wing 98, 95; tail
96, 94; bill 30, 28 mm.

“ Tris red, feet grey, maxilla black, mandible yellow.”

These two specimens as well as several others from Bogota are
undoubtedly different from J/. p. poliopis. They are much larger,
with a stouter, stronger bill; the forehead is broadly white,
bordered posteriorly by a very distinct, black band ; the cinnamon-
rufous colour below extends down over the breast, leaving only
the middle of the abdomen white, the flanks being mixed with
dull earthy brown, while in all the races of M/. panamensis the
breast and sides are sharply striped with blackish; the upper
parts are of quite another shade of brown, ete. The sexes are
alike, whereas there is a marked sexual difference in the forms
of M. panamensis.

M. mystacalis inhabits the mountains of Colombia, being found
in the Western as well asin the Eastern Cordillera (Bucaramanga,
Bogota).

172. MoNASA PALLESCENS Cass.

Monasa pallescens Cassin, Proc. Ac. N. Sci. Philad. 1860, p. 134
(1860.—Cordilleras of the Rio Truando) ; idem, |. c. 1864, p. 287,
pl. iv.; Wyatt, Ibis, 1871, p. 374 (Paturia, Magdalena Valley) ;
Sclater & Salvin, P. Z.S. 1879, p. 5386 (Remedios, Neche).

No. 2562. Q ad. Juntas, Rio Tamana (405 ft.), 5.111.09.—
Wing 142; tail 132; bill 39 mm.

“Tris dark brown, feet black, bill scarlet.”

A fine example of this scarce species, which is at once known,
among the other white-fronted forms, by its black throat and
smaller, nearly white upper wing-coverts. It corresponds exactly
with Cassin’s description and figure.

1198 MR. C. E. HELLMAYR ON TILE

M. pallescens is peculiar to the forests of Western Colombia,
Juntas being the most southerly locality on record. Specimens
from the Central Cordillera (Remedios, Neche, Paturia) I have
not seen, but from Sclater’s remarks they would seem to be
somewhat different.

173. Capito QUINTICOLOR Elliot.

Capito quinticolor Elliot, Nouv. Arch. Mus. Paris, i. Bull.
p. 76, pl. iv. fig. 1 (1865.—“ Nouvelle Grenade,” Triana coll.) ;
Dalmas, Bull. Soc. Zool. France, xxv. 1900, p. 176 (¢ 9: El
Paillon, near Buenaventura).

No. 2689. gad. ‘Tadé, 230 ft., 2.vi.09.—Wing 82; tail 453;
bill 21 mm.

“ Trides reddish brown, feet dark grey, maxilla black, mandible
blue-grey.”

This bird agrees well with the figure of the type except
in having a distinct orange tinge along the middle of the lower
breast and abdomen. It is an adult male in perfect plumage.
C’. quinticolor is the rarest of the Barbets of South America, only
four specimens being on record. The type, in the Paris Museum,
was brought by Mons. Triana from Colombia, without any
further locality. Count Dalmas recorded a couple obtained by
Eugéne André near Buenaventura, province of Chocé, viz., in the
same general district whence our specimen also comes.

C. quinticolor is apparently confined to the humid, forest-
covered lowlands of Western Colombia.

174. Capito MACULICORONATUS Lawr.

Capito maculicoronatus Lawrence, Ann. Lye. N. H. N. Y. vu.
p. 300 (Jan. 1861.—Lion Hill, Panama); Sclater & Salvin,
P.Z.8. 1879, p. 537 (Remedios, Neche: Antioquia).

Nos. 2297, 2312. § gd ad. Noévita: 16, 19.xi.08.— Wing 88, 85 ;
tail 56, 54; bill 213 mm.

No. 2227. g§ nearlyad. Rio Cajén: 31.x.08.—Wing 85; tail
56; bill 22 mm.

Nos. 2298, 2313. 9 9 ad. Noévita: 16, 19.xi.08.—Wing 84;
tail 53, 54; bill 212, 22 mm.

No. 2028. 9 imm. Noanama: 28.vili.08.—Wing 78; tail 54;
bill 21 mm.

“Tris black (¢), dark brown (2), feet blue, bill blue, tip
black.”

Compared with two specimens from Panama in the Munich
Museum, the males have the yellow tinge across the lower
breast rather paler, but in other respects they are similar.

C. maculicoronatus ranges from Veragua and Panama south-
wards to Western Colombia, and inhabits exclusively the hot,
forest-clad lowlands and foot-hills.

BIRDS OF WESTERN COLOMBIA. 1199

In Western Ecuador, its place is taken by C. squamatus Salv.*,
which differs, in both sexes, in lacking the scarlet patch on the
flanks, in possessing a Jarge white patch on the outer web of the
tertiaries, and in having the forehead orange-red instead of
black. The female, too, has the feathers of the back and the
upper wing-coverts narrowly edged with white. The hitherto
undescribed male has—lke that of C. maculicoronatws—the
cheeks, throat and foreneck white, the chest light saffron-yellow,
and the upper parts uniform glossy black, but there are hardly
any black streaks on the flanks. J have examined a good series
from Carondelet (province HEsmeraldas) and Santo Domingo,
W. Ecuador.

175. CAPITO BOURCIERII SALVINI Shelley.

[| Micropogon Bourcierit Lafresnaye, Rev. Zool. viii. p. 179
(1845—ad Bogotam ; deser. ¢).|

Capito salvint Shelley, Cat. B. Brit. Mus. xix. p. 119 (1891.—
“Central America, from Panama to Costa Rica,” no type
specified !); Dalmas, Bull. Soc. Zool. France, xxv. 1900, p. 180,
note (Las Cruces, Western Cordillera above Buenaventura,
6000 ft.). ;

C. bourciert (nec Lafresnaye) Sclater & Salvin, P.Z.S. 1879,
p- 538 (Frontino, Antioquia).

No. 2806. g ad. Loma Hermosa, Rio Jamaraya, 4150 ft.,
23.x.09.— Wing 74; tail 52; bill 20 mm.

INoseZmoo 250 G Quads. Pueblo) Rice, 52005 fhe sda.
25.x.09.— Wing 76, 73; tail 52; bill 20, 193 mm.

Nos. 2739, —. 2 ad.,? imm. Pueblo Rico: 7.ix., Nov. 09.—
Wing 74; tail 50, 51; bill 20, 19 mm.

Nos. 2808, 3774. 9 Q ad. Loma Hermosa: 23.x.09; Siato,
5200 ft., 25.1x.09.— Wing 71; tail 49, 47; bill 19 mm.

“Tris dark red, feet greyish green (¢) or blue-grey (2), bill
greenish yellow or yellow.”

As already pointed out by Count Dalmas (J.c.) the birds from
W. Colombia are referable to C. 6. salvini. Compared with
a good number from Costa Rica and Chiriqui the specimens from
Chocé are slightly larger on the average, but in coloration they are
nearly similar. The males have the throat dark crimson which
gradually passes into the deep orange of the chest, while, in the
females, the broad black frontal band is immediately followed by
the golden olive of the crown, without the least trace of the pale
blue cross-band, so conspicuous a feature in C. 6. bowrcierit 2, of
the Hastern Cordillera (Bogoti).

Count Dalmas (/.¢. pp. 179-180) most correctly explained the
distinguishing characters of the three races of this group, and I
quite agree with him that the West Ecuadorian birds constitute

* This, 1876, p. 494, pl. xiv. (= 9; Santa Rita); Goodfellow, Ibis, 1902, p. 218
(Santo Domingo).

1200 — MR. C. E. HELLMAYR ON THE

a quite distinct form, immediately recognizable by the crimson
of the throat and foreneck being abruptly contrasted with the
clear sulphur-yellow belly without any orange tinge. His name
CU’. shelleyi* is, however, antedated by C. equatorialis Salvad. &
Festa t by several months. Of this distinct form I have
examined, besides the type actually at Tring, two fine adult
males from Quito and Gualea, in the Munich Museum, and two
males in poor condition, from Nanegal and Esmeraldas, respectively,
in the British Museum collection. The original locality “‘ Rio
Napo” is erroneous, the type being a skin of the well-known
Quito form.
The range of the three races known to me is as follows :—

(a) C. 6. bourcieriti Lafy. Eastern Cordillera of Colombia
( Bogotz coll.), and Eastern Ecuador.

(b) C. 6. salvini Shelley. Costa Rica, Chiriqui and Western
Cordillera of Colombia.

(c) C. b. equatorialis Salvad. & Festa. Western Ecuador:
Intac, Chimbo, Pallatanga, Nanegal, Esmeraldas, ete.

[176. SEMNORNIS RAMPHASTINUS Jard.

Tetragonops ramphastinus Jardine, New Edinburgh Philos.
Journ. (n.s.) i. p. 404 (Oct. 1855.—‘ Hastern Cordillera between
Quito and the mountain Cayambe,” N. Ecuador).

No. 87. g ad. La Tigra, W. Cordillera, 5700 ft., 13.iv.,1899.
EH. André coll.— Wing 110; tail 81; bill 22 mm.

“Tris red, feet olive-yellow, bill. yellow, with a dusky spot
towards the tip.”

This bird agrees. perfectly with specimens in the Munich
Museum from the neighbourhood of Quito.

So far as I know the species has not previously been recorded
from Colombia. |

177. RamMpHastos swaAtnsonit Gould.
Rhamphastos Swainson Gould, P. Z.S. 1833, p. 69 (Sept.

1833—“ in montosis Columbie.”)

Ramphastos tocard (nec Vieillot) Sclater & Salvin, P. Z.S. 1879,
p. 537 (Concordia, Medellin, Remedios).

Rk. tocardus Cassin, Proc, Acad. N. Sci. Philad. 1860, p. 136
(River Nercua).

No. 2000. Q ad. Noanama {100 ft.), 22.viii.08.—Wing 230 ;
tail 170; bill 166 mm.

‘“‘Tris dark green; feet blue; base of maxilla below blackish,
diagonal line dark red, remainder yellow; mandible dark red,
apical third black with an oblong yellow patch near the tip.”

This bird is typical swainsonii, having the basal portion of both
mandibles (in skin) rich salmon colour, while in the allied

* Bull. Soc. Zool. France, xxv. p. 179 (Noy. 1900.—* Rio Napo ”’—errore !)
+ Boll. Mus. Zool. Torino, xy. no. 368, p. 22 (Feb. 1900.—Intac, W. Ecuador).

BIRDS OF WESTERN COLOMBIA. 1201

R. ambiguus Swains.* the same parts are dull black. C/r. Hartert’s
observations in Noy. Zool. v. 1898, p. 498.

I am at a loss to understand how the name R. focard Vieill.t
could have ever been applied to the present species. Levaillant’s
figure represents a bird with white throat and chest, and crimson
upper tail-coverts, whereas R. swainsoniz has the latter pure white
and the throat deep yellow. The next available name is the one
used above.

R. swainsonii (tocard auct.) ranges from southern Central
America through Colombia to Western Ecuador.

178. PrEROGLOSSUS ERYTHROPYGIUS SANGUINEUS Gould.

[Pteroglossus erythropygius Gould, P. Z. 8. 1843, p. 15 (July
1843.—Voyage of the ‘Sulphur’; the type locality, later given
as Realejo, Nicaragua, is doubtless erroneous). |

P. sanguineus Gould, Monogr. Ramph., 2nd edit., text to pl. 21
(1854—loe. ign.) ; Cassin, Proc. Acad. N. Sci. Philad. 1867, p. 109
(R. Truando, N. Colombia); Hartert, Nov. Zool. v. 1898,
p. 498 (Cachabi, Paramba, N. W. Ec uador).

P. erythropygius (nec Gould) Cassin, Proc. Acad. N. Sci. Philad.
1860, p. 136 (R. Truando).

No. 2022. gad. Noanama: 27.viii.08.—Wing 151; tail 175
bill (measured with chord) 130 mm.

Nos. 2023, 2024, 2163, 2189. 9 9 ad. Noanama: 27.viu.08 ;
Sipi: 6,14.x.08.—Wing 147-152; tail 170-182; bill 112-120
mm.

“Tris yellow; feet greyish green; maxilla greyish yellow, tip
yellow, broad culminal band and streak along cutting-edge black ;
mandible black.”

The series proves beyond doubt that P. e. sanguineus is a
perfectly valid form, the coloration of the bill being quite
constant in all five specimens. I have also examined ten skins
from various localities in N.W. Ecuador, belonging partly to the
Tring and partly to the Munich Museums.

These fifteen examples invariably have the lower mandible
black, while the maxilla, in addition to the black stripe along the
cutting-edge, shows a broad, black culminal stripe, extending over
more than the two basal thirds of its length. Cassin (/. e. pp. 109—
110) gives the same coloration for six specimens obtained on the
River Truando by Lieutenant Michler’s Expedition. Sometimes
the extreme base of the mandible is shightly clouded or marbled
with dull greyish yellow, and a small spot at the extreme tip is
soiled yellowish.

In P. e. erythropygius Gould, on the other hand, the bill is pale
yellow except a black stripe along the cutting-edges of the
maxilla, and a slight dusky tip to the lower mandible. The
culminal stripe is generally absent, though occasionally specimens

* Zool. Ilustr. iii. pl. 168 (no locality).
tone Dict. xxxiv. p. 281 (1819—based on Leyvaillant, Ois. de Paradis, etc., 11.
pl. 9).

1202 MR. GC. E. HELLMAYR ON THE

may be found with an indistinct, narrow streak down the centre
of the culmen.

P. é. sanguineus inhabits the lowlands and _hill-ranges of
Western Colombia (from the Truando southwards) and N.W.
Keuador (province Ksmeraldas : Cachabi, Paramba, Bulin, etc.)*.

P. ¢é. erythropygius replaces it in the more southern districts of
Western Ecuador: Babahoyo, Pallatanga, Chimbo, Rio Peripa,
Santa Rita, Santo Domingo, ete. The Tring and Munich Museums
possess good series of this form, showing its characters to be
quite constant f.

179. Coceyzus MELACORYPHUS Vieill.

Coccyzus melacoryphus Vieill. Nouv. Dict. viii. p. 271 (1817—
ex Azara: Paraguay).

No, 2104. g ad! Sip: /224%08— “Winey 110 tant 128)
bill 25 mm.

“Tris black, feet grey, bill black.”

Rather small, but not otherwise different from Brazilian
examples.

180. Pronopsirra puLcHRA Berl.

Pionopsitta pulchra Berlepsch, Ornith. Monatsber. v. p. 175
(1897.—San José, Rio Dagua, W. Colombia); Hartert, Nov.
Zool. v. 1898, p. 500 (Cachabi, N.W. Ecuador).

No, 2016. 2 ad. Noanama: 26.vii1.08.— Wing 146; tail 64;
bill (with chord) 22 mm.

“ Tris blue-grey, feet dirty yellow, bill white.”

A fine specimen of this rare Parrot, which was discovered
on the Rio Dagua by the late Gustav Hopke, and was afterwards
met with by Mr. Rosenberg near Cachabi, in the Hcuadorian
province of Esmeraldas.

The species is evidently restricted to the hot lowlands of
W. Colombia and N.W. Ecuador. It is allied to, but quite
distinct from, the Central American P. hamatotis Scl. & Salv.

181. Prionus mMEenstruvus Linn.

Psitiacus menstruus Linneus, Syst. Nat. 12, i. p. 148 (1766—
ex Edwards—hab. ign.—et Brisson: “ Guiane,” sc. Cayenne).

Pionus menstruus Sclater & Salvin, P. Z. 8. 1879, p. 538
(Remedios).

Nos. 2010, 2168, 2169. dg 6 @ ad. Noanama: 25.viii.; Rio
Garrapatas: 8.x.08.—Wing 180-175; tail 75-80; bill 27-

28 mm.

* Tawrence’s record of P. erythropygius from Chiriqui (Ann. Lye. N. H.N. Y. viii.
1865, p. 178) is certainly referable to some other species, perhaps P. torquatus, of
which the Munich Museum has a large series from Boquete.

+ Salvadori & Festa (Boll. Mus. Zool. Torino, xv. no. 368, 1900, p. 23) describe
a “young” bird of P. erythropygius (from Intac) as having the lower mandible and
a distinct culminal stripe black. I cannot help thinking that the specimen in

question really belongs to P. e. sanguineus, though its occurrence so far south would
be remarkable.

BIRDS OF WESTERN COLOMBIA. 1203

“Tris brown, feet whitish, maxilla black with base red, mandible
black.”

Colombian examples of this wide-spread species are apparently
not different from others taken in Cayenne, Brazil, and Venezuela.
There is perhaps generally more rosy suffusion on the foreneck,
though this is not quite constant.

182. MicrAsTUR GUERILLA INTERSTES Bangs.

| Micrastur guerilla Cassin, Proce. Acad. N. Sei. Philad. iv. p. 87
(1848.—Jalapa, Mexico). |

Micrastur interstes Bangs, Auk, xxiv. p. 289 (1907.—La
Estrella, Cartago, Costa Rica).

No. 2330. 9 ad. Noévita, Rio Tamandé (150 ft.), 24.xi.08.—
Wing 175; tail 165; bill (from cere measured with chord)
18 mm.

“Tris and feet yellow, bill black.”

This specimen, a perfectly adult bird in the “ plumbeous phase,”
except in being slightly larger, agrees with an adult from Miramar
(Chiriqui) and a couple from Cartago, Costa Rica (Underwood).
All the under surface, from the foreneck to the tail-coverts, is
closely barred black and white, as described by Mr. Bangs. The
sides of the head and the throat are pale smoky grey (the former
somewhat darker than the latter), exactly as in the Central
American skins; but the scapulars and, to a lesser degree, the
wings are more strongly tinged with chocolate-brown.

M. g. zonothorax Cab.* from Venezuela, of which I have
examined six specimens procured in the mountains around
Merida, differs from the Colombian and Costa Rican birds in
having the sides of the head and the throat dull rufescent brown,
and the quills more or less rufous-brown. Moreover, the mantle
is generally washed with chocolate-brown, whereby its close
relation to MW. rujficollis Vieill. is supported. That zonothorax
is merely a race of M. guerilla is clearly shown by a specimen
from Paramba, N.W. Ecuador, in the Tring Museum, which
combines the dull rufescent brown cheeks and throat with the
blackish wings.

M. g. interstes ranges from Costa Rica and Chiriqui southwards
to Western Kcuador (Nanegal, Paramba, Surupata, etc.).

183. ACCIPITER SUPERCILIOSUS Linn.
‘alco superciliosus Linneus, Syst. Nat. 12, i. p. 128 (1766.—

Surinam : juv.).

Accipiter tinus Sclater & Salvin, P. Z. 8. 1879, p. 541
(Remedios).

No. 2628. 2 imm. Tad6 (230 ft.), 30.iv.09.—Wing 149;
tail 105; bill 13 mm.

“Tris orange, feet yellow, bill black, base yellow.”

* Climacocercus zonothorax Cabanis, Journ, f. Orn, xiii, p. 406 (1865.—Puerto
Cabello, N. Venezuela).

1204 MR. CG. E. HELLMAYR ON THE

184, LEucoPreRNIS PLUMBEA Saly.

Leucopternis plumbea Salvin, Ibis (3) 1. p. 240, pl. vill. (1872.—
Ecuador); Hartert, Nov. Zool. ix. 1902, p. 605 (Paramba,
S. Javier, N.W. Ecuador).

No. 2103. g ad. Sipi, 150 ft., 29. ix.08.— Wing 220; tail 130;
bill 24 mm.

“Tris crimson, feet orange-red, bill black, base orange.”

This bird éoerontonee with Salvin’s description and figure with
the exception that the thighs are but obsoletely barred with
white. The other character istics, viz. the white under wing-
coverts and quill-lining, and the broad white bar across the middle
of the tail, are very well pronounced,

L. plumbea, although nearly allied to LZ. schistacea Sund, *
of Amazonia, may easily be distinguished by its much smaller size,
by lacking the white apical band £0 the rectrices, etc., ete. It was
hitherto known only as an inhabitant of Western Eeuador 7, and
is very rare in collections.

185. LeucoprerNis SEMIPLUMBEA Lawr.

Leucopternis semiplumbeus Lawrence, Ann. Lyc. N. H. N. Y.
vii. p. 288 (1861.—Panama Railroad).

LL. semiplumbea Sclater & Salvin, P. Z. 8. 1879, p. 540
(Remedios); Hartert, Nov. Zool. ix. 1902, p. 605 (Paramba, °
S. Javier, N.W. Ecuador).

No. 2566. 9 ad. Juntas, R. Tamand (405 ft.), 10.1ii1.09.—
Wing 188; tail 130; bill 22 mm.

“Tris yellow, feet orange, maxilla black, base orange, mandible
yellow.”

The skin is in every respect typical. L. semiplumbea ranges
from Nicaragua and Costa Rica through Western Colombia to the
province of Hsmeraldas, N.W. Ecuador.

186. HeERPETOTHERES CAGHINNANS Linn.

Falco cachinnans Linneus, Syst. Nat. x. p. 90 (1758—
ex Rolander: ‘‘ America meridionalis,” hab. subst. Surinam, auct.
Berlepsch, Nov. Zool. xv. p. 290).

Herpetotheres cachinnans Sclater & Salvin, P. Z. S. 1879,
p. 541 (Cauca, Remedios).

ING, ZO o AGl Vévita (150 ft.), 4.xii.08.—Wing 250;
tail 195; bill 26 mm.

‘Tris brown, feet yellow, bill black, base yellow.”

187. Lepropon PALLIATUS Temm.

Falco palliatus (Wied MS.) Temminck, Pl. Cel. livr. 23, pl. 204
(1823.—- “ Brésil et Guiane”; juv.).

* Asturina schistacea Sundevall, Ofvers. Vetensk.-Akad. Forhandl. vil. No. 5,
p- 132, note 3 (1850.—* Brasilia”).
+ Sharpe’s (Cat. B. i. p. 216) record from “ Panama” requires comfirmation.

BIRDS OF WESTERN COLOMBIA 1205

Leptodon cayennensis auct. *

No. 2721. 9 ad. Tado ea ft.), 26.vi.09.—Wing 270; tail 170
bill 29 mm.

“Tris dark grey, feet blue, maxilla black, mandible dark
blue.”

An adult bird of this wide-spread species.

188. CoLuMBA sPECcTOSA Gm.

Columba speciosa Gmelin, Syst. Nat. 1, ii 783 ete
Daubenton, Pl. Enl. 213: Cay enne) ; ches & Salvi Tit aA:
1879, p. 543 (Remedios).

No. 2476. g vixad. Noanama: 14.1.09.—Wing 187; tail 112;
bill 203 mm.

‘Tris and feet dark red, bill scarlet.”

A wide-ranging species.

189. CoLUMBA ALBILINEA ALBILINEA Bonap.

Columba albilinea (ew Gray MS.) Bonaparte, Consp Av. u
p- ol (Nov. 1854.—*“ Nova Granada ”’).

C’. albilineata Sclater & Salvin, P. Z. 8. 1879, p. 543 (Retiro).

No.2801. gad. Loma Hermosa, 4150 ft., 19.x.09.—Wing 195;
tail 140; bill 193 mm.

“Tris light brown, feet and bill lemon-yellow.”

This bird is exactly like specimens from Ecuador. North of
the Isthmus of Panama C. a. albilinea is represented by the
nearly allied C. albilinea crissalis Salvad. +, which has the under-
parts much paler and the top of the head more reddish.

190. CotumBA Goopsont Hart.

Columba goodsoni Hartert, Bull. B. O. C. xii. p. 42 (1902.—
5. Javier, Pambilar, and Carondelet, N.W. Ecuador) ; idem, Nov.
Zool. ix. 1902, p. 602 (Pambilar, 8. Javier, Carondelet, Rio
Sapayo t, Cayapas, N.W. Hcuador).

No. 2041. ¢ ad. Noanama (100 ft.), 31.viu.08.—Wing 150;
tail 113; bill 14 mm.

> tres pink, feet crimson, bill black.”

This specimen, a male in full plumage, agrees Rerieetly with two
topotypes from N.W. Heuador. As pointed out by Hartert, the
Species is a very distinct one and wey be distinguished inguin
C. nigrirostris brunneicauda Carriker $, of Costa Rica and Panama,
in having the top and sides of the head clear plumbeous grey and

* The specific name cayennensis cannot be used for this species, Falco
cayennensis Gmelin (Syst. Nat. 1, 1.1788, p. 269) being preoccupied by Falco
cayennensis of the same author (I. ¢. p. 263).

+ Columba crissalis Salvadori, Cat. B. Brit. Mus. xxi. p. 294 (1893.—Costa
Rica, etc.).

3p Erroneously spelt “ Japayo.”

§ Ann. Carnegie Mus. vi. Nos. 2-4, p. 395 (1910.—Guapiles, Costa Rica).

Prec. Zoot. Soc.—1911, No. LXX XI, dl

1206 MR. C. E. HELLMAYR ON THE

the throat light pearl-grey instead of vinous, the foreneck and
breast much paler, plumbeous washed with lilac, and the abdomen
dull vinous-brown. ‘The bright cinnamon-rufous colour of the
under wing-coverts and inner webs of the remiges serves to dis-
tinguish it at a glance from C. pluinbea bogotensis Berl. & Lev.
and C. subvinacea berlepscht Hart. *, found in the same districts.

C. goodsont is peculiar to the forest-covered lowlands (from sea-
level to about 500 feet) of N.W. Ecuador (province Hsmeraldas)
and Western Colombia.

191.: GEOTRYGON VERAGUENSIS CACHABIENSIS Hart.

[Geotrygon veraguensis Lawrence, Ann. Lyc. N. H. N. Y. viii.
p. 349 (1866.—Veragua). |

G. veraguensis cachabiensis Hartert, Nov. Zool. v. p. 504
(1898.—Cachabi, N.W. Ecuador).

Gi. v. cachaviensis Hartert, |. c. x. 1902, p. 603 (S. Javier, Rio
Sapayo, N.W. Ecuador ; crit..),

No. 2444. g ad. Noanama (100 ft.), 5.1.08—Wing 134;
tail 76; bill 182 mm.

‘“‘ Tris yellow, feet crimson, bill black.”

This specimen is decidedly darker sepia-brown on the back and
wings, and has a stronger violet-purple gloss on the upper mantle
than a single Costa Rican skin of true G. v. veraguensis. The
same differences were noticed by Hartert in a series from
N.W. Ecuador.

G. v. cachabiensis replaces the typical race in the Pacific
lowlands of W. Colombia and the adjoining parts of Ecuador
(province Esmeraldas).

192. GHorrRYGoN BOURCIERI Bonap.

Geotrygon bourcieri Bonaparte, Consp. Av. ii. p. 71 (Nov.
1854.— Lloa, Ecuador).

No. 2816. 2 ad. Pueblo Rico (5200 ft.), 27.x.09.—Wing 155;
tail 98; bill 17 mm.

“Tris yellow, feet light red, bill black.”

Compared with two fine adults from Hcuador (Aguapum) this
bird has the flanks slightly deeper rufous, and the forehead
less rosy, but these variations are very trifling.

This rare Pigeon was hitherto known only from Western
Ecuador (2 and N. Peru) and the present Hewat! extends its

range considerably to the north.

193. Rayncnortyx crnctus Salv.

Odontophorus cinctus Salvin, Ibis, (3) vi. p. 379 (1876.—Veragua;
= 6),

QO. spodiostethus Salvin, Ibis, (4) 1. p. 447 (1878.—Veragua ;
=o ye

* Noy. Zool. v. p. 504 (1908.—Paramba, N.W. Ecuador).

BIRDS OF WESTERN COLOMBIA. 1207

Rhynchoriyx cinctus Hartert, Nov. Zool. ix. 1902, p. 600 (Bultn,
Rio Bogot:i, Pambilar, N.W. Ecuador; crit.).

Nos. 2119, 2125. ¢ $§ ad. Sipi (150 ft.): 25, 28.1x.08.—
Wing 116, 120; tail 45; bill 17 mm.

No. 2126. 9 ad. Sipi: 28.1x.08.—Wing 116; tail 44;
bill 17 mm.

‘* Tris brown, feet blue, maxilla black, mandible grey.”

These specimens fully bear out the conclusions arrived at by
Hartert from the study of a series from N.W. Heuador, viz., that
O. cinctus and O. spodiostethus have been established upon phases
of one and the same species. The two skins, marked as “ ¢ ” by
the collector, agree substantially with the description ot
O. spodiostethus, having the sides of the head and throat bright
ferruginous, the chest dark cinereous, and the rest of the belly deep
ochraceous buff, the flanks finely vermiculated with dusky, and
the under tail-coverts only distinctly barred with black, ete.

The third specimen, sexed as “female,” tallies well with the
description of O. cinctus. It has the head above, nape, and chest
deep rufous-brown, while the remaining under paris are white,
broadly banded with black, except down the middle of the
abdomen; chin and upper throat are white; there is but an
indistinet, buffy whitish superciliary stripe, ete.

While there cannot be any further question as to O. cinctus
being the female, and O. spodiostethus the male of the same
species, yet the comparison of a good series from Veragua might
show the birds from the Pacific coast district of Colombia and
N.W. Heuador to be slightly different. In fact, the Sipi female
differs from Salvin’s and Grant’s descriptions in having the lower
back and rump bright rufescent brown, finely vermiculated with
dusky and spotted with black, especially down the centre, instead
of ‘‘dark grey or vinaceous, mottled with whitish.”

R. cinctus, wherever it occurs, appears to be arare bird. It has
been recorded from the Escondido River (Nicaragua), Panama,
Veragua, the Choco district (W. Colombia), and from the province
of Esmeraldas (N.W. Ecuador).

194. CHAMAPETES GOUDOTII GouDOTII Less.
Ortalida Goudotii Lesson, Man. d’Ornith. i. p. 217 (1828—

“les montagnes du Quindiu,” Colombia).

Chamepetes goudoti Sclater & Salvin, P. Z. 8. 1879, p. 544
(Retiro).

No. 2795. $ ad. Tatamdé Mountain, 4600 ft., 14.x.09.—
Wing 255; tail 245; bill 33 mm.

*“ Tris crimson, feet vermilion, bill black, bare space round the
eye dark blue.”

Identical with a specimen from Bogota. Birds from Ambato,
E. Keuador, are much brighter rufous underneath. They are
certainly subspecifically distinct and may belong to C. g. ruji-
ventris Tsch., from Peru, which I have not yet met with.

1208 MR, C. BE. HELLMAYR ON THE

195. NorHocercus INTERCEDENS Salvad..

Nothocercus intercedens Salvadori, Cat. B. Brit. Mus. xxvii.
p- 513 (1895.—Frontino, W. Colombia).

NV. bonapartii (nec Gray) Sclater & Salvin, P. Z.S. 1879, p. 548
(Frontino, Concordia).

No.2728. Qad. Pueblo Rico (5200 ft.), 16.viii.09.— Wing 203;
tail 70; bill 32 mm.

“Tris dark brown, feet grey, maxilla black, mandible grey.”

The specimen agrees with Salvadori’s description. The throat
is white, washed with pale rufescent buff on its lower portion.
The rump. upper tail-coverts, wings, as well as the abdomen, are
marked with numerous white or buffish dots.

NV. intercedens is peculiar to the Western Cordillera of Colombia
(Frontino, Concordia, Pueblo Rico). In southern Central America
it is represented by WV. frantzii Lawr.*, in the astern Cordillera
and in the mountains of Western Venezuela by 1. bonapartet
Gray ‘.

196. CRECISCUS ALBIGULARIS Lawr.

Corethrura albigularis Lawrence, Ann. Lyc. N. H. N.Y. vi.
p- 302 (Jan. 1861.—Panama Railroad).

Porzana albigularis Sclater & Salvin, P. Z. 5. 1879, p. 546
(Remedios).

Creciscus albigularis Hartert, Nov. Zool. ix. 1902, p. 604
(S. Javier, N.W. Ecuador).

No. 2144. $ vixad. Sipi, Rio Sipi, 1.x.08.—Wing 75; tail 30;
bill 18 mm.

“ Tris orange-yellow, feet dark greenish brown, bill black.”

This bird agrees with typical Panama specimens, the head above
being russet-brown, the chin and upper throat conspicuously
white, etc. There are a few cinnamon-rufous spots, but no white
bars, on the upper wing-coverts.

©. albigularis ranges from Panama south to N.W. Hcuador
(San Javier, province Esmeraldas).

197. JONORNIS MARTINICA Linn.

Fulica martinicu Linneeus, Syst. Nat. 12, i. p. 259 (1766—* in
Martinice inundatis ”).

Porphyrio martinicus Sclater & Salvin, 1. ¢. p. 546 (Medellin).

No. 2713. @ ad. Tadé (230 ft.), 18.v1.09.

“Tris light brown, feet yellowish brown, bill red, tip yellow.

9

198. ARAmIDESs wort Berl. & Tacz.
Aramides wolfi Berlepsch & Taczanowski, P. Z. 8. 1883, p. 576

®* Tinamus frantzii Lawrence, Ann. Lyc. N. H.N. Y. ix. p. 140 (1868.—Cervantes,
Costa Riea).

+ _Tinamus bonapartei G. R. Gray, List Spec, B. Brit. Mus., y. Gallinz,
p. 97 (1867.—Valley of Aragua, W. Venezuela).

BIRDS OF WESTERN COLOMBIA. 1209

(1884.—Chimbo, S.W. Ecuador); Salvadori & Festa, Boll.
Mus. Torino, xv. no. 368, p. 40 (Rio Peripa, W. Kcuador):
Hartert, Nov. Zool. ix. 1902, p. 604 (Pambilar, Carondelet,
N.W. Ecuador).

No. 2264. 9 ad. Névita: 11.xi.08.—Wing 160; tail 70; bill
55 mm.

“Tris scarlet, feet crimson, bill green, tip paler.”

This skin agrees perfectly with others from N.W. Ecuador,
notably with an adult female from Carondelet. Salvadori and
Festa (J. ¢.) have alveady pointed out that the description in the
Cat. B. Brit. Mus. xxiii. p. 55, is not quite exact.

A. wolfi is nearest to A. mangle Spix * and agrees with it in
the clear cinereous colour of the head and neck above, and in the
whitish throat, but may at once be distinguished by its much
larger bill, deep olivaceous-brown back, this colour passing into
rufous-brown on the mantle, black (instead of greyish) lower
abdomen and thighs, much darker, ruddy brown breast and upper
belly, ete.

A. wolfi is new to the fauna of Colombia. It was only reported
as an inhabitant of Western Keuador.

199. DENDROCYGNA DIscoLoR Scl. & Saly.

Dendrocygna discolor Sclater & Salvin, Nomencl. Av. Neotrop.
p- 161 (1873.— Venezuela, Guiana, et Brasilia,” type from
Surinam, cfr. Cat. B. Brit. Mus. xxvii. p. 162).

D. autwmnatis (nec Linneus) Cassin, Proc. Acad. N, Sci. Philad.
1860, p. 197 (R. Truando).

No. 2193. 2 ad. Mouth of Rio Sipi, near Noanama, 16.x.
08.—-Wing 225; tail 62; bill 50 mm.—“ Iris black, feet and bill
blood-red.”

Typical of D. discolor, the lower hind neck being buffy grey,
abruptly contrasted with the chestnut-brown back.

D. discolor is widely distributed in South America south of the
Panama Isthmus.

200. QUERQUEDULA CYANOPTERA Vieill.

Anas cyanoptera Vieillot, Nouv. Dict. v. p. 104 (1816—
“‘ yiviére de La Plata et 4 Buenos Aires”).

No. 2150. 2 ad. Sipi: 3.x.08.—Wing 175; tail 70; bill
40 mm.

“Tris black, feet putty brown, maxilla blue-grey, mandible ~
flesh-coloured.”

201. Carso vicua Vieill.
Hydrocorax vigua Vieillot, Nouv. Dict. viii. p. 90 (1817—ex
Azara: Paraguay).

* Gallinula mangle Spix, Av. Bras. ii. p. 74, pl. 97 (1825,—Brazil).

1210 MR. C. E. HELLMAYR ON THE

Carbo brasilianus? Cassin, Proc. Acad. N. &ci. Philad. 1860,
p: 197 (R. Truando).
~ No. 2045. 9 imm. Noanama: 1.ix.08.—“ Iris emerald-green,
feet black, maxilla grey, mandible yellow.”

IV. Conclusions.

Although our knowledge of Western Colombia is still very far
from complete, yet the researches of the naturalists mentioned
in the first part furnish sufficient evidence on which to base
some conclusions as to the general character and affinities of the
avifauna of this remote district. As might be expected from its
geographical situation, the fauna of Western Colombia shows a
mixed character, being composed partly of Ecuadorian and partly of
Central American species, the former, however, by far preponder-
ating. The most interesting result derived from Mr. Palmer's
collection is evidence of close similarity of the avifauna of the San
Juan Valley to that of the province of Esmeraldas, N.W. Ecuador.
In fact quite a number of remarkable species are apparently peculiar
to S.W. Colombia and the adjoining parts of N. Ecuador, while in
the more southern districts of Western Keuador other more or
less allied forms take their place. In several instances, however,
the Chocé region has a species or subspecies of its own, while
the province of Esmeraldas is inhabited by a representative form
generally distributed over Western Ecuador.

A few species range from Panama to N.W. Heuador, but do
not go further south. Several others extend from Panama to the
Dagua district, whereas in N.W. Ecuador another species or race
is found.

The subjoined tabular lists + may serve to illustrate these facts,
which are of some importance to the student of zoogeographical
problems.

(A) Species peculiar to Western Colombia (Chocé district) and
N.W. Ecuador (province Esmeraldas).

Turdus tristis dague. Dendrornis lachrymosa rostrata,
Henicorhina inornata. Cercomacra berlepschi.

Calospiza johanne. *Caprimulgus rosenbergi.

Hlenia cinerea parambe. Polyerata rosenbergi.
Rhynchocyelus cinereiceps flavotectus. *Hucephala hwmboldti.

Mionectes olivaceus hederaceus. Pionopsitta pulchra.

Chloropipo holochlora lite. Columba goodsont.

Sapayoa enigma. Geotrygon veraguensis cachabiensis.
Carpodectes hopkei. Leucopternis plumbea.

Automolus nigricauda. etc., etc.

+ The species not represented in Mr. Palmer’s collection, but procured by previous
travellers, are marked with an asterisk (*).

BIRDS OF WESTE

N COLOMBIA. IAL

(B) Species peculiar to Western Colombia.

Heleodytes albobrunneus.
Thryophilus nigricapillus schottii.
Huphonia xanthogaster chocoensis.
i. f. fulvicrissa.

Chlorochrysa nitidissima.
Calospiza palmeri.

Buthraupis melanochlamys.

B. aureocincta.

*Ostinops salmoni.
Masius chrysopterus bellus.
Siptornis erythrops griseigularis.
Thripadectes sclateri.
Anoplops bicolor dague.
Pittasoma rosenbergi.
Phethornis yaruqui sancti-johannis.
Thalurania fannyi.

* Adelomyia cervina.
Cyanolesbia kingii subsp.
Bucco noaname.
Capito quinticolor.

C. maculicoronatus (north to Panama).

Monasa pallescens.

Chloronerpes rubiginosus gularis.
* Picumnus olivaceus granadensis.

Pteroglossus e. sanguineus

(north to the Truando, south to N.W.

Eeuador).
Odontophorus parambe baliolus.
Nothocercus intercedens.

Represented in W. Ecuador by

T. n. nigricapillus.

HH. x. xanthogaster (°)

LE. fulvicrissa purpurascens (N.W.).
C. pheenicotis.

B. rothschildi (N.W.).

B. edwardsi (also in §. Colom)ia:
Pasto).

O. atrocastaneus.

M. c. coronulatus.

S. e. erythrops.

A. b. equatorialis.

P. rufopileatwn (N.W.).
P.y. yaruqui.

T. verticeps.

A. melanogenys maculata.
C. k. celestis.

|

C.

squamatus.

P. 0. harterti.
P. erythropugius erythropygius
(C. & S.W. Ecuador).

O. parambe paranbe.

(C) Species ranging from Panama to N.W. Ecuador, but not

known to occur elsewhere.

Chiromacheris vitellina.

Pachyrhamphus dorsalis.

Bucco pectoralis.

B. tectus subtectus.

(D) Species ranging from Panama to the Chocd district,

W. Colombia.

Cotinga nattererii.

Capito maculicoronatus.

1212

MR. C. E, HELLMAYR ON THE

(1) Species of Western Colombia represented in southern Central
America (Panama, Costa Rica) by nearly allied forms.

Western Colombia.

*Turdus tristis daque.

Heleodytes albobrunneus harterti.
TLeucolepis p. pheocephalus.

Thryophilus nigricapillus schottii.
+L. leucopogon.

* Henicorhina inornata.
+ Basileuterus t. tristriatus.
*Dacnis venusta fuliginata.

D. cayana corebicolor.
+Sporophila ophthalnica.
*Calospiza lavinia lavinia.

C. gyroloides gyroloides.

*O. johanne.
*O. larvata fanny.

Buthraupis melanoch lamys.

* Heterospingus wanthopygius.
*Hemithraupis salmoni.
}+Cacicus uropygialis.

Cyanocorax affinis affinis.

*Rhynchocyclus cinereiceps flavotectus.

+ Mionectes olivaceus hederaccus.

Myiobius sulphureipygius villosus.
+Pipra mentalis minor.

Chiromacheris vitellina.
+Tityra semifasciata columbiana.
*Lathria wnirufa castaneotincta.
*Lipaugus holerythrus rosenbergi.
*Carpodectes hopkei.

Siptornis erythrops griseiqularis.
+Hyloctistes subulatus assimilis.
+Xenops genibarbis littoralis.

* Dendrornis lachrymosa rostrata.
TD. triangularis equatorialis.
+Uyrmotherula fulviventris viduata.
+ Formicivora quixensis consobrina.
+Ramphocenus c. cinereiventris.
+Myrmelastes exsul maculifer.

Anoplops hicolor dague.
}Hormicarius analis destructus.

Pittasoma rosenbergi.
+Threnetes ruckeri fraseri.
*Chalybura urochrysa.

Chloronerpes rubiginosus gularis.

Picumnus olivaceus granadensis.
}Malacoptila panamensis poliopis.

Central America.

T. tristis cnephosa.

Hi. a. albobrunneus.

Lup. lawrencii.

T. semibadius.

T. thoracicus.

HL, leucosticta prostheleuca.
B. tristriatus melanotis.
D. v. venusta.

D. c. ultramarina.

S. aurita.

C. lavinia dalnasi.
tC. gyroloides bangsi.

C. florida florida and C. florida arcei.
C. larvata subsp.

B. arcei.

H.. rubrirfons.

HL. chrysomelas.

C. microrhynchus.

C. a. zeledoni.

R. c. cinereiceps.

MM. o. olivaceus.

IM. s. aureatus.

P. mentalis ignifera.

C. aurantiaca.

T. s. costaricensis.

Lu. clara.

Lh. holerythrus.

C. nitidus.

S. e. rufigenis.

A, s. virgatus.

X. g. mexicanus.

D. 1. lachrymosa.

D. t. punctigula.

M. f. fulviventris.

FE. q. boucardi.

R. c. semitorquatus.

MM. e. exsul.

A. b. bicolor.

EF. a. nigricapillus (Kastern Costa

Rica).

P. michleri (Panama).

T. r. ruckeri.

C. isawre (Veragua).

C. 1. uropygialis. :
P. 0. panamensis (Panama).
M. p. panamensis.

~ When also oceurring in the province of Esmeraldas (N.W. Ecuador), they are
marked with an asterisk (*): when more generally distributed in Western Ecuador,

with a dagger (t).

BIRDS OF WESTERN COLOMBIA. UALS

Western Colombia (cont.). Central America (cont.).
Aulacorhomphus albivitta pheolemus§. A. ceruleigularis.
*Pionopsitta pulchra. P. hematotis.
*Columba goodsoni. C. nigrirostris brunneicauda.
*Geotrygon veraguensis cachabiensis. G. v. veraguensis.
Nothocercus intercedens. NV. frantzii.

The evidence at hand is hardly sufficient to allow an exact
comparison of the fauna of Western Colombia with that of the
Central Cordillera, yet from what we know it appears that the
Pacific lowlands and foot-hills possess a considerable number of
peculiar elements, e. g. Supayou enigma, Cnipodectes subbrunneus,
Carpodectes hopkei, etc. Some of the species recorded only from
the Western Cordillera may turn up in the Central Cordillera ;
but, on the other hand, it is quite certain that a good many forms
that inhabit the central mountain-chain are altogether absent
from the Pacific slopes.

A thorough exploration of Southern and Central Colombia
cannot be too warmly recommended to the attention of
ornithologists, as offering a wide field for the solution of various
zoogeographical problems.

§ Aulacorhamphus pheolemus Gould, Ann. Mag. N.H. (4) xiv. p. 184 (1874.—
Concordia, Western Cordillera of Colombia (and Merida, in Venezuela—errore !)).

A. petax Bangs, Proc. Biol. Soc. Wash. xxi. p. 158 (1908.—San Antonio, R. Cali,
Western Cordillera of Colombia).

There can be no doubt that the blue-throated Awlacorhamphus, of the Western
Cordillera, has to bear the name A. a. pheolemus, which Mr. Bangs appears to have
overlooked. Although in the Cat. B. Brit. Mus. xix. p. 158, a specimen from
Merida, Venezuela, is given as type, this cannot be correct, for Gould (J. c.) clearly
descrined the Concordia bird (efr. “throat deep greyish blue”). It may be
mentioned that the Merida form was erroneously referred to pheolemus by Gould,
since a series in the Munich Museum undoubtedly belongs to A. a. albivitta, all the
examples having the throat white.

Proc. Zoou. Soc.—1911, No. LX XXII, 82

No. 100.

ABSTRACT OF THE PROCEEDINGS
oF THE
ZOOLOGICAL SOCIETY OF LONDON.*
October 24th, 1911.

Sir Joun Rose BrRAprForp, K.C.M.G., M.D., F.B.S8.,

Vice-President, in the Chair,

The Minutes of the last Scientific Meeting were confirmed.

The Sacrerary read a Report on the Additions that had been
made to the Society’s Menagerie during the months of June,
July, August, and September 1911.

Mr. JAMEs DunBAr-Brunton sent for exhibition two skins and
a mounted skull, with horns, of Bushbuck shot by him in North-
East Rhodesia.

Mr. W. B. Corton, of the Indian Civil Service, exhibited a
number of heads and horns of various species of Gazelles which
he had obtained in the Eastern Sudan, and gave a brief account
of their habits and distribution.

Mr. R. I. Pocock, F.R.S., F.L.S., F.Z.S., Superintendent of
the Gardens, exhibited two photographs of a living male specimen
of Red-fronted Gazelle (Gazella rujfifrons) which was presented to
the Society by Col. Julian Hasler in May 1908, and came from
Kano in Northern Nigeria. This specimen he proposed to make
the type of a new subspecies to be named G. rujifrons hasleri,
since it differed apparently from all previously recorded races of
that species in having the upper side of the nose pure white from
the nostrils back to the preorbital glands.

* This Abstract is published by the Society at its offices, Zoological Gardens,
Regent's Park, N.W., on the Tuesday following the date of Meeting to which
it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge,
to all Fellows who subscribe to the Publications ; but it may be obtained on the
day of publication at the price of Stapence, or, if desired, sent post-free for
the sum of Sia Shélings per annum, payable in adyance,

48

My, J. Lewis Bonnorrg, M.A., F.Z.S., exhibited two “ waltzing ”
Rats (Mus ratius) which he had bred in the course of his experi-
ments. They appeared in the F, generation, and it was noted —
that the strain had become weak and degenerate, other individuals
of that generation being born blind or otherwise defective. As
the “ waltzing” character had been proved to have a Mendelian
inheritance in Mice, it would be an interesting fact that, should
this character prove to have a Mendelian inheritance in Rats, it
would be a case of the genesis of a Mendelian character brought
about by artificial conditions, e.g. environment. ‘The original
stock were normal wild individuals. Mr. Bonhote also pointed
out that the two varieties of J. rattus found in Egypt, viz.
M.r. tectorwm with white underparts and IZ. r. alecandrinus with
dark underparts, had a Mendelian inheritance, the former white-
bellied form being dominant to the dark-bellied form. These
experiments also included the study of the inheritance of a fawn-
coloured variety, hitherto unknown, which appeared as a ‘‘ sport”
from wild-caught parents. ‘This variety had also a Mendelian
inheritance, the fawn-coloured ones being recessive to both the
normal wild forms.

Mr. D. Sera-Smuiru, F.Z.S., Curator of Birds, exhibited a
spirit-specimen of a nestling Australian Regent-Bird (Serzeulus
melinus) which had been hatched in the aviary of Mr. Reginald
Phillipps, of 26 Cromwell Grove, West Kensington, during the
past summer.

Mr. E.G. Boutencrr, Curator of Reptiles, gave the description
of anew Tree-Frog from Trinidad, living in the Society’s Gardens.
The Frog, which was brought back by Dr. Lewis H. Gough in
July last, was one of the smallest of the genus Hyla, and was

vemarkable for the rapid changes in colour and markings which
it displayed.

Mr. Bruce F. Cummines. read a paper, communicated. by
Mr. T. A. Cowarp, F.Z.S., on “ Distant Orientation in Batrachia,”
based on observations and experiments made by the author in
North Devon. ‘Two species of Newts had been used for the
experiments, and the results obtained lent support to the hypo-
thesis that these batrachians possessed a homing faculty, but no
very definite instinct for detecting water, even from a short
distance. Of the factors discussed in connection with amphibian
migration, it was suggested that in regard to Newts, a com-
bination of their homing faculty and their marked tendency to

walk downhill was chiefly of assistance to them in finding water
in which to breed.

Mr. OuprieLp Tomas, F.R.S., F.Z.8., read a paper on Mammals
collected in the Provinces of Sze-chwan and Yunnan, W. China,
by Mr. Malcolm Anderson, for the Duke of Bedford’s Exploration

49

of Kastern Asia. The paper formed No. XV. of the series, and

would be the last on Mr. Anderson’s specimens, as he was now
returning finally to America. During his work on the exploration

he had obtained 2700 specimens, besides many birds, and had
quite revolutionized our knowledge of the area explored.

The present collection, given, as before, to the National Museum
by the Society’s President, consisted of 160 specimens, belonging
to 33 species. ‘The following were described as new :—

RHYNCHONAX ANDERSON, gen. et sp. nn.

Allied to Uropsilus sor VS, but with p’ and i, present ‘no p,.
Head and body 70 mm.; tail 67; hind foot 15-5 5; skull 21-7.
Hab. Omi-san. Type. Male. No. 11.2.1.25. M.P.A. 2504.

NASILLUS GRACILIS, gen. et sp. nn.

Allied to the above, but with p* and p, present ; no i,.
Head and body 66 mm.; tail 55; hind foot 13°5; skull 20-5.
flab. Chin-fu-san. Type. Female. No. 2566.

SOREX WARDI FUMEOLUS, subsp. n.
Larger and darker-coloured than true wardi; the brain-case

broader.

Head and body 60 mm.; tail 60; hind foot 13; skull 18-1.
flab. Wei-choe, Si-ho R. Type. Male. No. 2627.

SORICULUS IRENE, sp. n.

Allied to macrurus, but brain-case lower.
Head and body 60 mm. ; tail 90; hind foot 16; skull 17-2.
Hab. Yuen-ching, Sze-chwan. Type. Female. No. 2673.

CHODSIGOA LARVARUM, sp. n.

Near Ch. hypsibia, but brain-case narrower and higher.

Head and body 68 mm. ; tail 50; hind foot 14; skull 18°8.
Hab. Imperial tombs HE. of Peking. TJype. Female. B.M.

No. 8.8.7.21.

APODEMUS SPECIOSUS ORESTES, subsp. n.

_ Size medium. Tail long; ears medium. Colour near sepia.
Head and body 93 mm.; tail 125; hind foot 24; ear 16.

Hab. Omi-san. Type. Male. B. M. No. 11.2.1. 170.

APODEMUS SPECIOSUS LATRONUM, subsp. n.

Size large. Tail short; ears long. Colour brown.
Head and body 107 mm. ; tail 101; hind foot 25; ear 20.
Hab. 'Ta-tsien-lu. Type. Male. B.M. No. 11.2.1.156.

MiIcROTUS MILLICENS, sp. n.

Skull flattened; an extra angle on m’*; tail long.

Head and body 90 mm. - tail 53 5 hind foot 18° 5 skull 24°3.
Hab. Wei-choe, Si-ho R. Type. Male. No. 2615.

50

Microrus (HoTHENOMYS) MELANOGASTER BLEUSIS, subsp. n.

Tail comparatively long; m° with four inner angles,

Head and body 98 mm.; tail 55; hind foot 17; skull 24-7.
Hab. BH. of Chao-tung-fu, N. Yunnan. ype. Male. No. 2696.

Microrus (EoTHENOMYS) OLITOR, sp. n.

Near MW. melanogaster, but no extra internal angle on m’.
Head and body 82 mm.; tail 34; hind foot 16; skull 24.
Tab. Chao-tung-fu. Zype. Female. No. 2714.

Microrus (CARYOMYS) ALCINOUS, sp. n.

Like Jf. (C.) eva, but colour very much darker.
Head and body 90 mm.; tail 56; hind foot 17; skull 24.
fab. Wei-choe, Si-ho R. Type. Male. No. 2631.

_ The Sscrerary presented a paper by Mr. EH. P. Srepsine,
E.LS., F.Z.S., entitled “‘ Game Sanctuaries and Game Protection
in India,” in which the author discussed the question of the
formation of Game Sanctuaries and what had been already done
in this direction in various parts of the country. Suggested
additions to the proposed New Indian Game Act were given,
and ‘close seasons’’ for certain species recommended as being
necessary for the preservation of the game of the country.

The next Meeting of the Society for Scientific Business will
be held on Tuesday, November 7th, 1911, at half-past Hight
o'clock P.M., when the following communications will be made :—

1. R. 1. Pococs, F.B.S., F.L.S., F.Z.S.
Lantern exhibition on the Moulting of the Arctic Fox.
fos SHES SINT Meas LEEUW

_ On the Moulting of the King Penguin (Aptenodytes pennanti)
in the Society’s Gardens. (With lantern illustrations.)

3. 7. E. Gunn, F.LS.

On the Presence of Two Ovaries in certain British Birds,
more especially the Faleonide. (With lantern illustrations.)

5l
4, Prof. P. P. Susuxin, C.M.Z.S.

Ontogenetical Transformations of the Bill in Ardea cinerea.

5. A. D. Imus, D.Sc., B.A.

On some Collembola from India, Burma, and Ceylon, with a
Catalogue of the Oriental Species of the Order.

The following papers have been received :—

1. H. B. Preston, F.Z.8.

Diagnoses of New Species of Terrestrial and Fluviatile
Shells from British and German East Africa.

bo

. R. LYDEKKER.
On the Milk-Dentition of the Ratel.

Structure of the Alimentary Canal of the Stick Insect
(Bacillus rosstvi Fabr.), with a Note on the Parthenogenesis of
this Species.

4. Ropert SHELFORD, M.A., B.Z.S.

Ginter, amongst the Blattide ; with a Revision of the
Genus Pr osoplecta Sauss,
5. The Rev. O. Pickarp-Campripen, F.R.S., C.M.Z.S8.
Contributions to the Knowledge of the Spiders and other
Arachnids of Switzerland.

A further Collection of Mammals from Egypt and Sinai.

7. J. T. Cunnincuam, M.A., F.ZS.

Mendelian Experiments on Fowls.

8. Prof. Grorrrey Surrn, M.A.

The Freshwater Crayfishes of Australia,

52

Communications intended for the Scientific Meetings should
be addressed to
P. CHALMERS MITCHELL,

Secretary.

ZOOLOGICAL Society oF Lonpon,
Recrnt’s Park, Lonpon, N.W.

October 31st, 1911.

No. 101.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.*
November 7th, 1911.

FrepDErick Ginuert, Esy., Vice-President, in the Chair.

The Minutes of the last Scientific Meeting were confirmed.

Mr. F. Menterra Ocitvis, F.Z.S., communicated a paper by
Mr. T. E. Gunn, F.L.S., ‘‘On the Presence of Two Ovaries in
certain British Birds, more especially the Falconide.” The
author outlined the views held by the majority of English mor-
phologists on the reproductive organs of adult female birds, and
enumerated examples which he had collected during a number of
years where the right as well as the left ovary was present, and,
so far as could be ascertained, in the two cases of which sections
had been made, where the right ovary was functional. He
pointed out the extraordinary preponderance in his examples of
paired ovaries occurring in the Falconide, as compared with those
derived from any other source, and remarked that in that family
the ovaries were usually placed symmetrically one on either side
of the vertebral column and at about the same level. In examples
other than the Falconide this symmetrical arrangement was the
exception rather than the rule, the right ovary generally occupy-
ing a position almost directly below the left, in the left half of
the body-cavity, which, in the author’s opinion, suggested a half-
way home on the road leading to the final disappearance of the
right ovary.

Mr. R. I. Pocock, F.RB.S., F.LS., F.Z.S., Superintendent
of the Gardens, exhibited a series of lantern-slides prepared from

* This Abstract is published by the Society at its offices, Zoological Gardens,
Regent's Park, N.W., on the Tuesday following the date of Meeting to which
it refers. It will be issued, along with the ‘Proceedings,’ free of extra charge,
to all Fellows who subscribe to the Publications ; but it may be obtained on the
day of publication at the price of Szarpence, or, if desired, sent post-free for
the sum of Six Shillings per annum, payable in adyance.

54

photographs taken by Mr. P. W. Farmborough, F.Z.S., illustrat-
ing the colour-change and phases in the moult of an Arctic Fox
now living in the Society’s Gardens,

Mr. D. Seru-Smirs, F.Z.S., Curator of Birds, read a paper,
illustrated by lantern-slides, on the Moulting of the King Penguin
(Aptenodytes pennanti) in the Society’s Gardens. He referred
to Mr. de Winton’s paper on the same subject which appeared
in the ‘ Proceedings’ in 1898. The specimen observed by
Mr. de Winton did not moult until it had lived sixteen months
in the Gardens, whereas the specimen now in the menagerie
had moulted twice in six months.

The author stated that the new feathers were almost fully
grown before the old ones were shed, and that the latter had to
be rubbed off by the bird’s beak or feet, as they were firmly
attached to the sheaths of the new feathers. The paper was
illustrated by photographs, which showed the bird in various
stages of the moult, as well as by specimens of the shed feathers.

Dr. S. F. Harmer, M.A., F.R.S., V.P.Z.S., read a paper by
Prof. A. D. Imus, D.Sc., B.A., entitled, ‘‘Some Collembola from
India, Burma, and Ceylon, with a Catalogue of the Oriental
Species of the Order.” Four genera and twenty-eight species
were described as new, amongst the latter the most remarkable
being a form unique among Collembola in possessing a median
cercus to the fifth abdominal segment, and for the reception of
which a new subfamily was formed. The total number of Col-
lembola known from the Oriental region was stated to amount to
53 species comprised within 27 genera.

A paper on the “‘ Ontogenetical Transformations of the Bill in
Ardea cinerea,” by Prof. P. P. Susuxin, C.M.Z.S., was read by
Mr. D.Sera-Smiru, F.Z.8. The author gave a description of the
gradual development of the bill in a series of embryos and young
specimens of the Heron upon which he had made observations.
The simple rhamphotheca proved to be only a late stage of the
compound one, and the form of the Ardeine bill he regarded as
a derivative one, and discussed its resemblance to those of allied
forms.

55

The next Meeting of the Society for Scientific Business will
be held on Tuesday, November 21st, 1911, at half-past Hight
o'clock P.m., when the following communications will be made :—

1. Dr. Grorrrey Suiru, M.A.

The Freshwater Crayfishes of Australia.

2. Frank E. Bepparp, M.A., F.R.S., F.Z.S8.

Contributions to the Anatomy and Systematic Arrangement
of the Cestoidea.—III. On a New Genus of Tapeworms from
the Bustard (Hupodotis kori).

(SU)

. ALFRED HE. Cameron, M.A., B.Sc.

Structure of the Alimentary Canal of the Stick Insect
(Bacillus ross Fabr.), with a Note on the Parthenogenesis of
this Species.

ne

. G. A. Boutencer, F.R.S., F.Z.S.

Some remarks on the Habits of British Frogs and Toads,
with reference to Mr. Cummings’s recent communication on
“« Distant Orientation in Amphibia.”

5. H. B. Preston, F.Z.S.

Diagnoses of New Species of Terrestrial and Fluviatile

Shells from British and German East Africa.
6. R. LyDEKKER.
On the Milk-Dentition of the Ratel.

The following papers have been received ;—~
1. Ropert SHetrorp, M.A., F.Z.S.

Mimicry amongst the Blattide; with a Revision of the
Genus Prosoplecta Sauss.

2. The Rev. O. PrckArp-CamsBrinax, F.R.S., C.M.Z.S.

Contributions to the Knowledge of the Spiders and other
Arachnids of Switzerland.

3. J. Lewis Bonunors, M.A., F.L.S., F.Z.8.

A further Collection of Mammals from Egypt and Sinai,

4. J. T. Cunnineuam, M.A., F.Z8.

Mendelian Experiments on Fowls.

56

Communications intended for the Scientific Meetings should
be addressed to

P. CHALMERS MITCHELL, |
Secretary.

ZooLocicaL Society or Lonpon,
ReceEnNtT’s Park, Lonpon, N.W.

November 14th, 1911.

No. 102.

ABSTRACT OF THE PROCHEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.*

November 21st, 1911.

S. F. Harmer, Esq., M.A., Sc.D., F.R.S., Vice-President,
in the Chair,

The Minutes of the last, Scientific Meeting were confirmed.

The Secretary read a Report on the Additions that had been
made to the Society’s Menagerie during the month of October
issih:

Mr. R. I. Pocock, F.R.S., F.Z.S., Curator of Mammals,
exhibited two living examples of an Elephant-Shrew (JJacro-
scelides sp.) from Bechuanaland, which had been presented to the
Society by Capt. H. O. F. Littledale and forwarded by Dr. L.
Péringuey, C.M.Z.S., and remarked that these specimens, together
with another in the same consignment, but belonging to a different
species, were apparently the first representatives of the genus
Macroscelides the Society had ever possessed. After drawing
attention to some interesting structural features shown by the
animals, Mr. Pocock said that the Society was to be congratulated
upon the safe arrival of these Elephant-Shrews, because, apart
from members of the Hedgehog-family, which from being omni-
vorous are more easy to preserve in captivity, exotic species of
Insectivora are proverbially difficult to keep alive for any length
of time and are therefore seldom exhibited in the Gardens,

Dr. Grorrrey Smiru, M.A., read a paper, communicated by
the SECRETARY, entitled “‘ The Freshwater Crayfishes of Australia.”
The object of this paper was to revise the classification and

—

* This Abstract is published by the Society at its offices, Zoological Gardens,
Regent’s Park, N.W., on the Tuesday following the date of Meeting to which
it refers.. It will be issued, along with the ‘ Proceedings,’ free of extra charge,
to all Fellows who subscribe to the Publications; but it may be obtained on the
day of publication at the price of Stxpence, or, if desired, sent post-free for
the sum of Stix Shillings per annum, payable in advance.

58

nomenclature of the Australian and Tasmanian Crayfishes, and to
give diagnoses of the genera and species with their limits of
distribution. The work was based on a large material obtained
from all parts of the continent and from Tasmania. Many of
the specimens had been collected by the author, but the majority
formed a very large collection brought together during the past
twenty years by Prof. Baldwin Spencer.

Four genera were recognised, Astacopsis, Cherops, Parache-
rops, and Hngeus: the first three genera included the Freshwater
Crayfishes proper, and the last-named genus contained the Land-
Crayfishes, which were not dealt with in this paper.

The geographical distribution of these genera and its bearing
upon the geographical problems of Australia was discussed, and
the view was supported that the Bassian Subregion represented
the home of the Australian Crayfish, and that Astacopsis was
nearest the ancestral form.

Mr. F. KE. Bepparp, M.A., F.R.S., F.Z.S., Prosector to the
Society, presented a paper on “A new Genus of Tapeworms from
the Bustard (Hupodotis kori).” Four complete specimens and
some fragments of this Cestode had been obtained from a
S. African Bustard in the Society’s Gardens, and the author
regarded it as a member of the group Tetracotylea, but could not
reconcile its characters with those of any other genus of that
group at present known. He briefly described its anatomical
characters and discussed its systematic position, and proposed a
new genus and species for its reception.

The SECRETARY presented a memoir by Mr. A. E. Cameron,
M.A., B.Sc., entitled “‘ The Structure of the Alimentary Canal of
the Stick-Insect, Bacillus rossii, Fabr., with a Note on the
Parthenogenesis of this Species.”

The author stated that this insect had a rather limited dis-
tribution, occurring in the south of Europe and in the north of
Africa, and that in the wild state it was not found north of
Orleans. Certain peculiarities of the alimentary canal were dealt:
with which were regarded as adaptations to the mode of life of
the species. Attention was drawn to the fact that the male was
only rarely found in the wild state, and that parthenogenetic
reproduction of B. rossii had been verified, for during four
generations the specimens kept by the author had showed no
males. The fact that the males were disappearing suggested that
parthenogenesis was not the primitive method of reproduction,
but that the species had become adapted to it through the failure
of sexual reproduction.

Mr. H. B. Preston, F.Z.S., communicated a paper based on a
collection of Terrestrial and Fluviatile Shells made by Mr. Robin
Kemp in British and German East Africa. One new genus and
thirty-four new species were described, which represented only a
very small portion of the large number of species collected

59

Mr. EH. G. Bounencer, F.Z.S., presented a short paper by
Mr. G. A. Boutencsr, F.R.S., F.Z.S., containing some remarks
on the habits of British Frogs and Toads, for the information of
those who might feel inclined to carry out further observations on
the subject of the migrations of Amphibia as dealt with in a paper
recently read before the Society. The Common Toad was strongly
recommended as the most suitable Batrachian on which to
institute series of experiments on Distant Orientation.

A paper on the “ Milk-Dentition of the Ratel” was received
from Mr. R. LyprexKsr, in which he described an instance of
primitive features present in the milk-dentition being entirely
lost in the teeth of the permanent series. So far as he was aware,
no such atavistic feature had been hitherto recorded in the case
of any existing mammals.

The next Meeting of the Society for Scientific Business will
be held on Tuesday, February 6th, 1912, at half-past Hight
o’clock p.M., when the following communications will be made :—

1. Mrs. R. Harte THomas, F.Z.S.
On Experimental Pheasant Breeding.

2. J. T. Cunninenam, M.A., F.Z.S.

Mendelian Experiments on Fowls.

3. H. G. Primer, F.R.S., F.L.S., F.Z.S.

Report on the Deaths which occurred in the Zoological
Gardens during 1911,

4, J. Lewis Bonnorte, M.A., F.L.S., F.Z.5.
A further Collection of Mammals from Egypt and Sinai.

The following papers have been received :—

1. Rosert SHetrorD, M.A., F.Z.S.

Mimicry amongst the Blattide; with a Revision of the
Genus Prosoplecta Sauss.

60

2. The Rev: O. Pickarp-Camprince#, F.R.S., C.M.Z.S.

Contributions to the Knowledge of the Spiders and other
_ Arachnids of Switzerland.

3. Herpert L. Hawnins, M.Sc., F.G.S.
The Classification, Morphology, and Evolution of the Echi-
noidea Holectypoida.

4. H. Lysrer Jameson, M.A., D.Sc., Ph.D., F.Z.S.

Studies on Pearl-Oysters and Pearls.—I. The Structure of
the Shell and Pearls of Margaritifera vulgaris Schumacher :
with an examination of the Cestode Theory of Pearl Pro-
duction. .

Communications intended for the Scientific Meetings should
be addressed to
P. CHALMERS MITCHELL,

Secretary.

ZOOLOGICAL Society oF Lonpon,
RecENt’s Park, Lonpon, N.W.
November 28th, 1911.

Papers (continued).

Page
40. On a new Species of Dinotherium (Dinotherium hobleyi) from British East Africa.

By C. W. Anprews, D.Se., F.R.S., F.Z.S. (British Museum, Natural History).

(Ge TE NGTS VATA) oops ain eet we eee anee Rar eeMeney FURS ABD AISS sw sow o:ni 2 4 cto heal cote ang OcheD
41. On an Amphipod from the Transvaal. By the Hon. Paun A. Mernuen, F.Z.8.

(O15. NUTONE STENT) aa sc otis fote te NS onan etal eA CREE Seg eles AVen NE eae’ a) a's.s een Sata : 948
42. An African Rhinoceros, Klipspringer, and Gazelle. By R. Lyprxker. (Text-figs.

AQT T G38) Wee Maar co age ics aptreretae te chances Lottie ay QUIRED EC 2c Oe aR eRe 958

43. The Subspecies of the Spanish Ibex. By Prof. Ange Caprmra, O.M.Z.S. (Pls. LIL—
DONE errata eset IS AI) Se oles ieee Sree Sra Sc cloit 32 e 35 oir eo OA ee SO EamMeMmE

44. On Antelopes of the Genera Madogua and Rhynchotragus found in Somaliland. By
R. HE. Draxe-Brockman, M.R.C.S., L.R.C.P., F.Z.S8. (Pls. LV. & LVI.) ........-- O77

45, Contributions to the Anatomy and Systematic Arrangement of the Cestoidea.
II. On. Two new Genera of Cestodes from Mammals. By Franx EB. Brppanp, >
M.A., F.R.S., F.Z.S., Prosector to the Society. (Text-figs. 204-215.) ............ 994

46. Some Madreporaria from the Persian Gulf. By Rurn Harrison, Oxford. Witha
Note on the Memoir and some Further Notes on Pyrophyllia inflata by Sypney J.
Hickson, M.A., D.Se., F.R.S., F.Z.S. (Pls. LVII. & LVIII. and Text-figs. 216-221.) 1018

47. On Variation in the Medusa of Merisia lyonst. By Cuaruns L. Boutencer, M.A.,

F.Z.8., Lecturer on Zoology in the University of Birmingham. (PI. LIX. and
Mexteie se 2oBe) ay .tevers -f lsrede iste ieletors SNE LiG Cod OCR TIG ROOD ASIC SP PEA aOR OOO Be, do 1045

48. The Marginal Processes of Lamellibranch Shells. By Cyrim Crosszanp, ¥.Z.8,
(Pl. LX. and Text-figs. 229 & 230.) ....-. RES CARH Cars Soar n ean clas 1057

49. Warning Coloration in a Nudibranch Molluse and in a Chameleon. By Cynrin
ROSSINI REZESS Sete Saya eer eraies ecperetaratelchsitae Cede SR aR tant BRE CGA O ONS 1062

50. Chromodorids from the Red Sea, collected and figured by Mr. Cyril Crossland. By

Sim Crags sbmom ake CAVE Gear C Be BZN. (RIM PCIEys erseietacc are selina tienes 1068
51. On some new South African Permian Reptiles. By R. Broom, D.Sc., C.M.Z.S.

Calg EO OUR MG UNI) SG so Sock eal TERA heed aN ec A ra ne ee Lr
52, On a new Tree-Free from Trinidad, living in the Society’s Gardens. By Epwarp G.

Bovunenerr, Curator of Reptiles. (Pl. LXIV.).......... eT a ii has Serccecrs ecru 1082
53. A Contribution to the Ornithology of Western Colombia, By ©. E. Hetumayr,

Division of Birds, Zoological Museum, Munich ............ SO EEOC REC One eb ae 1084
STIG ere Ia Gein tl (a8 Oboe She O San Oeenh cio seD USS ch Ociph vont Senos lane Bee ade eMW oc i
INE ole Camron each) oa ee cols gp vasaed ee eRe O canoe BA oboe AEA Sood Jeo il
List of Contents............. ener ess NEN shea hater aol iran gdh gcabde (eee Xe oth.s CR iti
Alphabetical List of Contributors ............-- Hei aceasta MaSteR NR ay aie Senay eas ix
News Genericul erinsirnts sielecisutihsavers (atta ater sloasrsi cia esehawa ce eet lesearalsy chen seeseie Seeteent Ne te XV1il
TndextomSerentiticwNames’ \voustrrtccncts acactor tena: sheer as chsh nr) <selipia oa pepc sce) pale ats ia xix

Index of Illustrations

Tiara Micmn eee a evateh niatectetoie roles XXXVI

LIST ‘OF PLATES. —

1911, Parr IV. (pp. 869-1213).

Plate Page
XLIII.
XLIV. Aleyonaria of the Cape of Good Hope ...........-..... - 870
XLV.
XLVI. Skulls of Cynodont Reptiles ................000.....0. 893
XLVII. Tooth-germs in Macropus billardierd ........0-..0ee+s000 926
SCUMLLL. emenhencu men ayant, saistctatcieisetciale\ a lacie eats 943
XULIX.
L. ! HLWwenGng ONY xe TOUTED. es sarees cin voi iale'vm «wie ele e tyem ss eel e : 948
LI.
LIL. Capra pyrenaica hispanica. (Summer Pelage.) .......-..
LI. Capra pyrenaica victorie, § 2. (Summer Pelage.) ...... |
LIV. Capra pyrenaica victorig. (Winter Pelage.) ............
LY. 1. Madoqua phiillipsi gubanensis. 2. Madoqua phillipsi
hararensis. 8. Madoqua phillipsi ......++ BSP oe OT7
LVI. 1. Madoqua piacentinti. 2. Madogqua swaynei .......+++
ae \ Madreporaria from the Persian Gulf .........+2+..--.. 1018
MINS Variation) Merisia Wyomst 6 ice. «sees ovine <2 <a\* =) </- 1045
LX. Marginal processes in Lamellibranch Shells.............. 1057
LXI. Chromodorids from the Red Sea ............00...-: -.. 1068
enna) Fossil Reptiles from South Africa,............. 0. 0eceee 1073
LXIII. J
LXIV. Gough’s Tree-Frog (Hyla gought) 0... ..eceescceccceenes 1082
NOTICE. .

The ‘ Proceedings’ for the year are issued in four parts, paged consecutively,
so that the complete reference is now P. Z. 8.1911, p.... The Distribution
is as follows :—

Part I. issued in March.

~~, Pp ical Ul Re eae June.

eae 3) Ee ate September.

ae me Wier December.
\) ‘ Proceedings,’ 1911, Part III. (pp. 557-868), were published on
aN" September 15th, 1911.
ea

‘The Abstracts of the Proceedings,’ Nos. 100-102, are
contained in this Part. “.

*p 970

i :
SY
au

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